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BIGNONIACEAE. Almost ninety species of this beautiful order are described by authors, the greater part of which are at present incorporated among the genuine species of Bignonia of Linne; a genus that will hereafter be divided, according to the shape of the calyx, the number of fertile stamina, and more especially the form of the fruit (which in some species is an orbicular or elliptical capsule, varying in others to a long cylindrical figure, with seeds partly cuneated, or thickened at one extremity, and in others, a truly compressed Siliqua) together with the relative position of the dissepiment, in respect to the valves of the fruit.
The greater portion of Bignoniaceae appears to exist in the equinoctial parts of America; Some, however, are natives of India, and a few occur on the western coast of Africa, and Island of Madagascar, but in Terra Australis the order is reduced to four plants, of which one is a recent discovery, and may be referred to Spathodea. In that continent, the order exists only upon the North and East Coasts; it is not, however, entirely limited to the tropic, for Tecoma of Mr. Brown is also found in latitude 34 degrees South, on which parallel it has been traced at least three hundred and fifty miles in the interior to the westward of the colony of Port Jackson.
ASCLEPIADEAE and APOCINEAE. Nearly the whole of the plants in the recently formed herbarium, that belong to these natural families, have been described from specimens formerly discovered upon the East and North Coasts, several of which appear to give a partial character to the vegetation of some parts of its shores.
Hoya (hardly Asclepias carnosa of Linne) Cynanchum, Gymnema, Gymnanthus, Sarcostemma, and probably Secamone, as belonging to Asclepiadeae, and all the genera of Mr. Brown (Lyonsia excepted) referred to the latter order, exist on that extensive coast, where Balfouria and Alyxia have each an accession of species. Of Strychnos, which is also frequent, and probably produces its flowers during the rainy season (as has been remarked of this genus in other countries) specimens in that stage of its fructification are still a desideratum; all that is known respecting the plant being the form and size of its fruit, which in some species varies considerably.
GOODENOVIAE. The Herbarium contains very few specimens of this considerable Australian family, the greater mass existing in and to the southward of the parallel of Port Jackson. The order is reduced to Goodenia, Scaevola, Velleia, and the tropical Calogyne on the North-west Coast, and the few species of the two first genera prove to have been formerly discovered upon the South Coast during the voyage of Captain Flinders, of which one plant has alsa a much more extensive range than has been given it heretofore. It is Scaevola spinescens, which forms a portion of the harsh, rigid vegetables of Dirk Hartog's Island on the West Coast, and from that shore probably occupies a part of a very considerable extent of barren country in the interior, in a direction towards the East Coast, having been seen in abundance in the latitude of Port Jackson, so near that colony as the meridian of 146 degrees 30 minutes East. A new Velleia, discovered on the North-west Coast in latitude 16 degrees, augments that genus, belonging to the section with a pentaphyllous calyx.
RUBIACEAE. The existence of several plants of this extensive family in the intratropical parts of Terra Australis especially when aided by some individuals of almost wholly exotic tribes, that form a prominent feature in the Flora of other equinoctial countries, tend, in some measure, to diminish the peculiar character of the vegetation of Terra Australis on those shores, and thus it is a considerable assimilation to the Flora of a part of a neighbouring continent that has been traced. About thirty species are preserved in the collections of these voyages, for the most part belonging to genera existing in India, but more abundant in the tropical parts of South America.
Of these, Gardenia, Guettarda, Cephaelis, Coffea, Psychotria, and Morinda, are found on the East Coast; whilst, in corresponding parallels on the opposite, or north-western shores, the order, although not materially reduced, is limited to the two latter genera, with Rondeletia, Ixora, and Genipa.
It is worthy of remark, that the range of Psychotria, which has not been observed beyond the tropics in other countries, extends in New South Wales as far south as the latitude of 35 degrees; at the western extremity of which it does not appear to exist.
CAPRIFOLIAE, Juss. The situation of Loranthus and Visvum, in the system, appears to be undetermined by authors. M. Jussieu associated them with Rhizophora, in the second section of this order, from which Mr. Brown has separated this latter genus, and with two others found in Terra Australis, has constructed a distinct family, named Rhizophoreae; suggesting, at the same time, the analogy of Loranthus and Viscum to Santalaceae, and particularly to Proteaceae. The genus Loranthus, of which nearly the whole of its described species have been limited to the tropics, is, however, sparingly scattered on all the Coasts of Australia, where about eleven species have been recently observed, parasitical chiefly upon certain trees that constitute the mass of the forests of that vast continent; namely, Eucalyptus, Casuarina, Acacia, and Melaleuca.
A solitary and very remarkable deviation from the usual natural economy of Loranthus, is observed in a species (L. floribunda) described and figured by M. Labillardiere, which is found on the shores of King George's Sound, where, in no way recognising the dependent habits of its congeners, it rises from the soil to a tree fifteen feet high, being never remarked relying upon other vegetables for its subsistence. Viscum is found in the colony of Port Jackson, to which it is not confined, having been also gathered at Endeavour River, on the same coast, within the tropic. The southern range of the two genera seems to be nearly beyond the fortieth degree of latitude; but in the northern hemisphere, Loranthus exists in Siberia.
UMBELLIFERAE. The equinoctial portion of the Herbarium contains only three or four plants of this extensive European order, belonging to Hydrocotyle, Azorella of Cavanilles and Labillardiere (from which Trachymene of Rudge is probably not distinct) and a suffruticose plant referred to Cussonia, that have been collected upon the East Coast. Upon the north-western shores, Azorella was alone remarked, of which a species is very general upon its main and islands, and chiefly remarkable for its gigantic herbaceous growth.
MYRTACEAE. With respect to that portion of Myrtaceae, lately discovered upon the north-western shores of Australia, and which are alone worthy of remark here, it is to be observed, that, considering the many points of that coast visited during the progress of the relative voyages, the number of species observed are comparatively few, for, including Eucalyptus, it does not exceed sixteen plants. Of Eucalyptus itself, only seven species were detected on those shores, and these, for the most part, form small trees, more approaching the average dimensions of all their congeners in the colony of Port Jackson. Melaleuca is limited to three species, one of which was originally discovered by the celebrated navigator, Dampier, on the West Coast, where Beaufortia has been recently seen. Four species of Tristania, their related genus, were gathered in about latitude 15 degrees South, where also an Eugenia, bearing fruit, was observed; but of Leptospermum, or Baeckea, genera chiefly belonging to the higher latitudes of New Holland, no species appeared throughout the whole extent of coast examined.
RHAMNEAE and CELASTRINAE were formerly united among the Rhamni of Jussieu, but disposed in sections, differing from each other in the position of the stamina, with relation to the petals, and in the character of the fruit; which, when viewed with other important differences of fructification, induced Mr. Brown to modify and define them as distinct orders.
In the Herbarium of the voyages, there are a few plants belonging to Rhamnus, Ziziphus, Ceanothus, or Pomaderris, and Celastrus, but both families prove to be comparatively rare in the intratropical parts of Terra Australis, beyond which Cryptandra seems only to exist. Upon the north-western shores, a species of Ziziphus (common to the East and North Coasts) forms a tree of large dimensions, where also an undescribed Celastrus has been discovered. Since Pomaderris evidently increases from the verge of the tropic southerly towards the parallel of Port Jackson, where its maximum exists, and as it is frequent on the South Coast, it is highly probable the West Coast is not wanting of the genus, particularly as traces of it were found on Dirk Hartog's Island.
LEGUMINOSEAE. There are upwards of one hundred and forty species of this extensive natural class in the Herbarium recently formed, which bear a proportion to the aggregate of the entire collections of about one to nine.
Of the Australian portion of Mimoseae, which (having been met with upon all the coasts of the continent, and equally diffused in the interior) forms a leading characteristic of its vegetation, upwards of fifty species have been collected, in various stages of fructification; nearly the whole of which are unpublished plants. Several of those discovered on the north-western shores, and islands off the West Coast, being also extremely curious in their general form and habits; and the existence of a few appears limited to a solitary particular situation, and no one species was observed common to those parts, and the opposite or eastern shores of the continent.
The Papilionaceous division exceeds seventy species, two-thirds of which belong to established diadelphous genera, found chiefly within the tropic, where some, peculiar to Terra Australis, and heretofore limited to the more temperate regions, have been discovered. Thus Hovea and Bossiaea were detected in New South Wales, in latitude 20 and 22 degrees South, as well as on the North Coast; the latter genus being likewise found on the north-western shores, where also two species of Kennedia exist; and Templetonia, a genus nearly related to Bossiaea, originally discovered on the southern shores of Australia, is abundant on an island off the West Coast.
Upon the North-west Coast, particularly in the parallels of 14 and 15 degrees South, where an exotic feature (if the usual characteristic of the Flora of other countries might in this case be so termed) is as manifest, and is as strongly blended with the pure Australian character (Eucalyptus and Acacia) in its general vegetation, as on any other parts of those shores; Jacksonia and Gompholobium, genera of Papilionaceae, with distinct stamens, almost limited to the parallel of Port Jackson and the South Coast, were observed: Daviesia, almost wholly restricted to the higher Australian latitudes, has been remarked on the North Coast. Of Lomentaceae, Bauhinia, Caesalpinia, and the emigrant genus Guilandina, are all of intratropical existence in New South Wales, as also upon the North-west Coast; but Cassia, although it has an equal extensive range in the equinoctial parts of New Holland, has also been recently traced as far in the interior, on the parallel of Port Jackson, as the meridian of 146 degrees East.
EUPHORBIACEAE. The Herbarium contains thirty-three plants of this very numerous order, whose maximum seems decidedly to exist in India and equinoctial America. The whole of the Australian species are referable to established Linnean genera, of which Croton and Phyllanthus are most remarkable and numerous, existing on all the intratropical shores of Terra Australis, but by no means limited to them, both genera, together with Euphorbia and Jatropha, being found in the parallel of Port Jackson; and Croton exists likewise at the southern extreme of Van Diemen's Land, which is probably the limit of the genus on that hemisphere.
A Tragia (scarcely distinct from a species indigenous in India) is sparingly scattered on the East and North Coasts; and Acalypha has been remarked on these, as well as the north-western shores.
PITTOSPOREAE. Of this small family, whose characters and limits were first described by Mr. Brown, there are sixteen species in the Herbarium of these voyages, referable to Bursaria, Billardiera, Pittosporum, and two unpublished genera.
Billardiera, whose species are wholly volubilous, and which are not found north of the parallel of Port Jackson, is frequent on the South-west Coast, and has been recently remarked on the West Coast of Van Diemen's Land. Bursaria on the other hand, appearing limited to New South Wales, has been traced within the tropic to latitude 19 degrees South on those eastern shores, and although the genus Pittosporum is even more extensively diffused on that coast, it has not been met with upon the north-western shores, whilst the islands off the West Coast furnished me with two new species.
DIOSMEAE, although very frequent in the higher latitudes of Terra Australis, where they are so frequent as to give a peculiar character to their vegetable productions, is comparatively rare within the tropic; for upon the East Coast Eriostemon and Phebalium appear to be the only genera, the latter having been recently discovered, in about latitude 20 degrees South.
With some undescribed species of Boronia, a new genus allied to Eriostemon has been observed on the north-western shores, in the parallel of 15 degrees South, having a remarkable pinnatified fimbriated calyx.
Of the related family ZYGOPHYLLEAE (an order proposed by Mr. Brown to be separated from the Rutaceae of Jussieu) Tribulus is frequent on the tropical shores of New Holland, and a species of Zygophyllum, with linear conjugate leaves and tetrapterous fruit, was remarked upon an island off Shark's Bay, on the West Coast.
MELIACEAE. The several genera of this order, whose maximum is in the equinoctial parts of America, differ from each other in the form of the remarkable cylindrical nectarium, the situation or insertion of the antherae upon it, as well as the character of its almost wholly capsular fruit. This structure of nectarium is most striking in Turraea, of which a species was observed upon the East Coast, far within the tropic; where also, as well as on all the other equinoctial shores of the continent, Carapa, more remarkable on account of the valvular character of its capsules, and the magnitude and irregular figure of its nuts, is very general, and probably not distinct from the plant (C. moluccensis, Lam.) of Rumphius, who has given us a figure in his Herbarium Amboinense volume 3 table 61, 62.
SAPINDACEAE. Of the very few plants referred to the family in the Herbarium, two genera are only worthy of remark here, the one an Ornitrophe, found on the East Coast, in about latitude 35 degrees, as also within the tropic; and the other, which appears to belong to Stadmannia, was discovered upon the same coast, in latitude 31 degrees South, the type of the genus being the bois de fer of the French colonists, a timber tree indigenous at the Island of Mauritius.
MALVACEAE, Juss. Tiliaceae, Juss. Sterculiaceae, Vent. Buttnericeae, Brown. These several families, of which the first is by far the most extensive, have been viewed by Mr. Brown, as so many allied orders of one natural class, to which the general title of Malvaceae might be applied. About thirty-six species of these orders collectively, are preserved in the present Herbarium, referable at least to eleven genera, of which nine are most abundant in (and form a characteristic feature of) the botany of India, and the equinoctial parts of South America. Fourteen species of Hibiscus and Sida were observed on the intratropical Coasts of Australia, beyond which also, on the opposite shores of the continent, each genus has been remarked. One species of Bombax with polyandrous flowers, and subspherical obtusely pentagonal capsules, was discovered upon the East Coast, in about latitude 14 degrees South, and on nearly the western extreme of the same parallel, it appeared much more abundant. Of Sterculia which is scarcely to be found beyond the tropics in other countries, a species exists in New South Wales in the latitude of 34 degrees, on which parallel it is more frequent in the western interior, and in that direction it has been traced to the distance of three hundred miles from the sea-coast. The genus is also found on the North and North-west Coasts, where the species assume more particularly the habits of their congeners in India. Among the plants of this family in the Herbarium is a species of Helicteris (as the genus stands at present) which was observed on the North-west Coast bearing fruit, wanting the contortion that characterizes the genus.
This plant, together with three other described species, having straight capsules, may hereafter be separated from that Linnean genus, and constitute a new one of themselves. Grewia, Corchorus, Triumfetta, and Waltheria, have been observed upon the North-west Coast, where also Abroma, hitherto limited to the tropical parts of New South Wales, has been discovered bearing flowers and young fruit. One species of Commersonia was gathered at widely-different parts of the north-western shores, and Lasiopetalum, whose species are more general at both extremes of the parallel of the colony of Port Jackson, has been also seen just within the tropic on the East Coast, and at Dirk Hartog's Island, off Shark's Bay, on the opposite shore.
CAPPARIDES. At least ten species of Capparis have been discovered upon the coasts of Terra Australis, for the most part within the tropic, but of these the fructification of two are wanting. A few have been detected on the East Coast, but they are more frequent and various in their species upon the north-western shores of the continent. Within an area on this extensive coast, not exceeding four degrees of longitude, on the parallel of 15 degrees South, a tree of very remarkable growth and habit, has been traced, having all the external form and bulk of Adansonia of the western shores of Africa. At the respective period of visiting those parts of the North-west Coast, this gouty tree had previously cast its foliage of the preceding year, which is of quinary insertion, but it bore ripe fruit, which is a large elliptical pedicellated unilocalar capsule (a bacca corticosa) containing many seeds enveloped in a dry pithy substance. Its flowers, however, have never been discovered, but from the characters of the fruit, it was (upon discovery) referred to this natural family. M. Du Petit Thouars has formed a new genus of Capparis pauduriformis of Lamarck, a plant of the Island of Mauritius, which he has named Calyptranthus. It has one division of the calyx so formed, that by its arcuated concavity (before expansion) it conceals the whole flower, and the other portions of the calyx; and should this genus be adopted by future botanists, a second species has been recently discovered upon Dirk Hartog's Island, although of remarkably different habit.
Cleome has been observed only in the equinoctial parts of Australia, and like Capparis, several species exist on the North-west Coast, being limited to C. viscosa in New South Wales.
Drosera, which Jussieu associates with these genera is generally diffused, being found within the tropic, at Endeavour River, and on the North-west Coast; at Port Jackson, and at the southern extremes of Van Diemen's Land.
DILLENIACEAE. To that Australian portion of the order lately enumerated by M. Decandolle, the present Herbarium offers, in addition, only two species of the genus Hemistemma of M. Du Petit Thouars. The one discovered on the North-west Coast, and allied to H. angustifolium of Mr. Brown; the other proving also new, but approaching in character the doubtful species, H. leschenaultii of Decandolle, and was discovered upon Rottnest Island, off the western coast of the continent, and is the first certain species of the genus, that is not limited to a tropical existence.
In addition to what has been advanced in respect to certain natural orders that appear in the Herbarium, formed under the stated circumstances, a slight mention might be made of other detached genera, or families sparingly observed on these coasts, that were more particularly investigated during the progress of the late voyages; but as these several plants form portions of orders so extremely limited, and in themselves presenting nothing remarkable in their internal structure, or external habit, a few remarks on a general comparison of the vegetation of the North-west Coast, with the other shores of Terra Australis, will conclude this notice.
It is very necessary to premise, that the plants observed and collected upon the North-west Coast, during the late voyages, are not to be considered as even a distant approach to an entire Flora of that extensive line of shore; since the long-established droughts of the seasons (as already remarked) in which the greater part of that coast was visited, had wholly destroyed plants of annual duration, with most of the Gramineae, and had indeed generally affected the mass of its herbaceous vegetation. The collections, therefore, can simply be viewed as a gleaning, affording such general outlines of characteristic feature, as will enable the botanist to trace its affinity to the more minutely defined vegetation of the other equinoctial shores of the continent, as well as perceive its general, and, in some instances, almost total want of relation to the botany of other parts, in the more temperate or higher latitudes, where certain striking peculiarities of the Australian Flora more particularly exist.
Upon a general comparison of those collections that were thus formed on the North-west Coast, with the plants of the North and East Coasts, aided also by some few observations made during the voyages, it appears that (with the exception of Gompholobium, Boronia, Kennedia, and one or two unpublished species not referred to any family) the genera (of which several are proper to India) are the same, although the species are very distinct upon the several coasts.
Notwithstanding an identity of genera has been remarked upon their opposite shores, there are, nevertheless, certain others, frequent upon the East Coast, that appear wholly wanting on the north-western shores: of these, the existence of some, even in the tropical parts of New South Wales, seems governed by the primary formation of the coast, its mountainous structure, and consequent permanency of moisture in a greater or less degree; namely, almost all the genera of Filices, the parasitical Orchideae, Piper, Dracontium and Calladium (genera of Aroideae) Commelina and Aneilema, Calamus and Seaforthia, Hellenia a solitary Australian genus of Scitamineae, some genera of Rubiaceae, particularly Psychotria and Coffea, certain genera of Asphodeleae, as Cordyline, and a genus allied to it, whose fructification is at length obtained, a solitary plant of Melastomeae, and an individual Nymphea.
Other genera also, but little influenced by those local circumstances of situation on the East Coast, that are excluded from the opposite shores, are Leucopogon (the only equinoctial genus of Epacrideae observed during the late voyages) the families Bignoniaceae, Jasmineae, the genus Erythrina, and of Coniferae, Araucaria of Norfolk Island. This absence of several orders of plants on the north-western shores, existing in New South Wales, or opposite coast, as well as the consideration (at the same time) of the evident causes of such a disparity of species on the former coast, would suggest the opinion, that such plants alone of other parts of the continent are indigenous to the North-west Coast, as are capable of sustaining themselves in a soil subjected to seasons of protracted parching droughts. This may apply to some species upon that coast, but it cannot be reduced to a general conclusion; for, on the one hand, it is singular so few of the plants of the South and South-west Coasts, and particularly that none other of their genera of Proteaceae (than those already mentioned) found altogether in an arid soil, should have been discovered throughout any part of its extensive shore; whilst, on the other hand, at a peculiar structure of a small and limited portion of that coast, in the vicinity of York Sound, a sufficiency of shade was observed to be actually produced by the unusually broken character of the country, to favour the nourishment and growth of certain plants alone to be seen beneath the shade of dense forests. These species were Myristica insipida, discovered by Mr. Brown, on one of the Prince of Wales group of islands on the North Coast; Cryptocarya triplinervis, Brown; bearing ripe fruit, Abroma fastuosa; and an undescribed Eugenia.
Although the several genera of plants lately observed on the north-western shores are also frequent in other equinoctial parts of the continent, there is, among the many species which are absolutely proper to that coast, a Capparis of such extraordinary habit, as to form a feature in the landscape of a limited extent of its shores, in the enormous bulk of its stem and general ramification, bearing a striking analogy to the Adansonia of the west coast of Africa.
The results of such observations on the vegetation as could only be made in a general way, at parts approaching each extreme of the North-west Coast, show their little affinity to each other; for the northern extremity partakes more fully of that feature of the line of coast contiguous to it, which (as already remarked) extends along the north-western shores, declines materially at, and in the vicinity of their southern limits, where the characteristic vegetation of the south, and perhaps the west, coasts has more particularly been found. Besides Eucalyptus and Acacia, which are abundant on every shore, and generally diffused throughout those parts of the interior that have been penetrated, there is another genus almost equally dispersed, which is, however, on the North-west Coast reduced to three species. This is Dodonaea, whose maximum is certainly in New South Wales, within and beyond the tropic, upon the coast, and generally in the interior of the country, extending also to the southern extremity of Van Diemen's Land.
Our very limited knowledge of the Flora of this vast continent (excepting of a part east of longitude 144 degrees, and included between the parallels of 31 and 35 degrees in New South Wales) is entirely confined to the vegetation of its immediate shores, upon every distinct coast of which, landings, more or less frequent, and under various circumstances, have been effected; although of all, very considerable portions remain unexplored, and of the line of West Coast (properly so denominated) the shores of Shark's Bay, and some few parts south of it, have alone been scientifically investigated. The interior within the tropic remains entirely in obscurity; the continental defect of a want of large streams having a distant source, to aid a penetration to the internal parts of the country, together with other effectual obstacles, draw at present a veil, and forbid all research into its Natural History and character, which will not be removed for very considerable periods (perhaps ages) yet to come!
It was the general remark made during a former expedition in the interior of New South Wales, that no absolutely entire change takes place in the vegetation east of the meridian of the new settlement named Bathurst; but that the plants of the coast were more or less frequent at a hundred and fifty miles from the sea, although in a country estimated at about two thousand feet above its level. Having to this circumstance added a remarkable and obvious sameness (arising from an extensive dispersion) of a vein of vegetation in a large tract of country, it may be inquired, how far these facts might, when applied to other parallels, identify a certain portion of the Flora of the interior, and that of the sea-coast in the same latitude; or, in other terms, how far the botany of the coast indicates the general feature of the vegetation to a certain limit, in the interior on the same parallel? Favourable opportunities were afforded me, to compare the vegetation of opposite coasts within the tropic, at the eastern and western extremes of a particular parallel; and the results of such a comparison identified many species on the two coasts. I have annexed a list of those plants that are common to the North-west and East Coasts in and about the parallel of 15 degrees South, from a contemplation of which, together with the above remarks, and a further comparison of the species with those of the shores of the Gulf of Carpentaria, through which that degree of latitude passes, might not a general idea of some portion of the Flora of the expanse of intermediate interior (far beyond the reach of actual investigation) be presumed?
A few observations relative to the geographical range of certain genera and species, hitherto considerably circumscribed, will close this notice.
The genus Pandanus has ever been viewed by botanists as equinoctial; nor was it till recently ascertained satisfactorily, that one of its species (P. pedunculatus, Brown) exists on the shores of Port Macquarie in New South Wales, in latitude 31 degrees South: and I have been credibly informed, that the same plant is frequent in the vicinity of Port Stephens, which is at least a degree to the southward of the above parallel. The latitude of 32 degrees South may be considered the utmost extreme of ranges from the equator of the genus in Terra Australis, on the opposite shore of which, as also in all other countries, it has not been remarked beyond the tropics.
The palms of Terra Australis, which (as previously observed) are remarkably limited on the north-western shores, have a very considerable diffusion on the North and East Coasts, and have even a more general dispersion on the latter shores, than has been allowed them formerly. Seaforthia is frequent in dense forests on the East Coast, almost to latitude 35 degrees South, where it exhibits all the tropical habits assumed on the northern shores, although the difference of climate, and consequent temperature, are abundantly obvious. On the other hand, a palm of very robust growth, with large flabelliform fronds, and spinous foot-stalks, was remarked at the head of Liverpool River, in latitude 12 degrees South, on the North Coast; and although without fructification, no doubt existed of its being the Corypha australis, hitherto limited to the shores and vicinity of Port Jackson.
Araucaria excelsa. The Norfolk Island pine, which, without doubt, must have been particularly noticed by the celebrated circumnavigator Captain Cook, in 1770, on the discovery of New South Wales, although the circumstance of the very general existence of a pine upon the islands and main of that coast, north of the Percy Isles, does not appear to be mentioned in the accounts of that particular voyage, has a far more extensive range upon that shore than has been hitherto understood. During the Mermaid's voyages, Araucaria was observed in the vicinity of Mount Warning, in New South Wales, which lies in the parallel of Norfolk Island (29 degrees South); thence northerly it was very sparingly seen towards the tropic, within which, however, as far as latitude 14 degrees, it is very abundant, forming upon several islands the only timber. This is probably the nearest approach of the species to the equinoctial line; and although it occupies an area of nine hundred miles, it is very probably limited in Terra Australis to its immediate shores; and, as appears to be the case with Pandanus, exists only within the influence of the sea air.
Calladium macrorhizon, Willd., formerly observed by Sir Joseph Banks, at Endeavour River, on the East Coast, has been recently detected in moist woods, in the country off which the Five Islands are situate, extending on that shore to latitude 35 degrees South: and Schelhammera multiflora, Br., a delicate plant of Melanthaceae, discovered likewise at Endeavour River, abounds in shady forests, in latitude 31 degrees, upon the same extensive coast.
The following plants, formerly considered as indigenous only in Van Diemen's Land, have been recently ascertained to exist also in New South Wales, in or about the parallel of the colony of Port Jackson.
Croton viscosum, Labill., originally discovered on the South-west Coast, was seen in the interior, as far to the westward of the colony as longitude 146 degrees East.
Croton quadripartitum, Labill., was observed in longitude 148 degrees.
Goodia latifolia, Salisb., was remarked sparingly in the interior, in the meridian of 147 degrees 30 minutes East: and Daviesia latifolia of Mr. Brown is very frequent in societies upon plains at Bathurst, in longitude 149 degrees East, where also Eryngium vesiculosum, of Labillardiere, was observed.
Aster argophyllus and obovatus, Labill. These two species were described by Mons. Labillardiere, from specimens gathered in the southern extremes of the above island, and have been lately seen tolerably frequent in a remarkable tract of country, in latitude 34 degrees, on the limit of the colony, where the former assumes a robust, arborescent habit. Aster phlogopappus, of the same eminent author, was recently remarked upon the more elevated parts of the Blue Mountain Range, on the margin of a remarkable cataract.
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A LIST OF PLANTS COMMON TO THE EAST AND NORTH-WEST COASTS OF TERRA AUSTRALIS, IN AND ABOUT THE PARALLEL OF FIFTEEN DEGREES SOUTH, WHERE THE BREADTH OF CONTINENT EXCEEDS 1800 MILES.
Gleichenia Hermanni, Br. Eriocaulon fistulosum, Br. Philydrum lanuginosum, Gaertn. Flagellaria indica, L. Dioscorea bulbifera, L. *? Pandanus pedunculatus, Br. Cycas angulata, Br. Santalum oblongatum, Br. Exocarpus latifolia, Br. Persoonia falcata, Br. Grevillea mimosoides, Br. Hakea arborescens, Br. Buchnera ramosissima, Br. Adenosma coerulea, Br. Orthostemon erectum, Br. Tabernaemontana orientalis, Br. Carissa ovata, Br. Strychnos lucida, Br. Alyxia obtusifolia, Br. Ipomoea longifiora, Br. Ipomoea denticulata, Br. Ipomoea maritima, Br. Evolvulus villosus, R. et Pav. Cuscuta carinata, Br. Cordia orientalis, Br. * Clerodendrum inerme, Br. * Avicennia tomentosa, L. Chionanthus axillaris, Br. Olea paniculata, Br. Maba laurina, Br. Sersalisia obovata, Br. Mimusops parvifolia, Br. Terminalia, sp. allied to Catappa, Lam. Cleome viscosa, L. Capparis sepiaria, L. Hibiscus tiliaceus, L. Abroma fastuosa, Br. Bombax australis. Jacksonia thesioides. Bauhiniae sp. Caesalpiniae sp. Cassia occidentalis, L. Guilandina Bonduc, L. Morinda citrifolia, L. * Carapa moluccensis, Lam. Zizyphus melastomoides. * Bruguiera gymnorhiza, Lam. Casuarina equisetifolia, Lam.
Should the botany of the shores of the Gulf of Carpentaria, in the vicinity of those parts, through which the above parallels pass, generally correspond (on comparison) with the above list, it is more than probable that these several species occupy portions of the intermediate interior bounded by the meridians of 125 and 145 degrees East; those plants excepted, having an asterisk prefixed to them, which as forming mangroves, or from other causes exist only on the sea shore.
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A LIST OF PLANTS OBSERVED DURING THE LATE VOYAGES ON THE SHORES OF TERRA AUSTRALIS, THAT ARE ALSO COMMON TO INDIA OR SOUTH AMERICA.
Acrostichum alcicorne, Sw. Polypodium acrostichoides, Sw. Nephrodium exaltatum, Br. Nephrodium unitum, Br. Vittaria elongata, Sw. Asplenium nidus, L. Daval1ia flaccida, Br. Gleichenia Hermanni, Br. Flagellaria indica, L. Dioscorea bulbifera, L. Calladium ? macrorhizon, Willd. Aristolochia indica, L. Daphne indica, L. Salicornia indica, Willd. Deeringia celosioides, Br. Plumbago zeylanica, L. Dischidia nummularifolia, Br. Acanthus ilicifolius, L. Acanthus ebracteatus, L. Ipomea Turpethum, Br. Ipomea denticulata, Br. Ipomea maritima, Br. Evolvulus villosus, R. et Pav. Trichodesma zeylanica, Br. Tournefortia argentea, L. Cordia orientalis, Br. Plectranthus scutellarioides, Br. Clerodendrum inerme, Br. Vitex ovata, L. Vitex trifolia, L. Avicennia tomentosa, L. Mimusops kauki, L. Aegiceras fragrans, C. Koenig. Scaevola koenigii, Vahl. Cleome viscosa, L. Capparis sepiaria, L. ? Calophyllum inophyllum, L. Morinda citrifolia, L. Carapa moluccensis, Lam. Sophora tomentosa, L. Cassia occidentalis, L. Guilandina bonduc, L. Abrus precatorius, L. ? Acacia scandens, Willd. ? Hibiscus tiliaceus, L. Suriana maritima, Jacqu. Pemphis acida, Forst. Rhizophora mangle, L. ? Bruguiera gymnorhiza, Lam. Sonneratia acida, L. Abroma fastuosa, Br. Casuarina equisetifolia, Forst.
...
CHARACTER AND DESCRIPTION OF KINGIA, A NEW GENUS OF PLANTS FOUND ON THE SOUTH-WEST COAST OF NEW HOLLAND: WITH OBSERVATIONS ON THE STRUCTURE OF ITS UNIMPREGNATED OVULUM; AND ON THE FEMALE FLOWER OF CYCADEAE AND CONIFERAE.
BY ROBERT BROWN, ESQUIRE, F.R.S.S.L. AND E. F.L.S.
(READ BEFORE THE LINNEAN SOCIETY OF LONDON, NOVEMBER 1 AND 15, 1825.)
In the Botanical Appendix to the Voyage to Terra Australis, I have mentioned a plant of very remarkable appearance, observed in the year 1801, near the shores of King George the Third's Sound, in Mr. Westall's view of which, published in Captain Flinders' Narrative, it is introduced.
The plant in question was then found with only the imperfect remains of fructification: I judged of its affinities, therefore, merely from its habit, and as in this respect it entirely agrees with Xanthorrhoea, included the short notice given of it in my remarks on Asphodeleae, to which that genus was referred.* Mr. Cunningham, the botanist attached to Captain King's voyages, who examined the plant in the same place of growth, in February, 1818, and in December, 1821, was not more fortunate than myself. Captain King, however, in his last visit to King George's Sound, in November, 1822, observed it with ripe seeds: and at length Mr. William Baxter, whose attention I had particularly directed to this plant, found it, on the shores of the same port in 1823, both in flower and fruit. To this zealous collector, and to his liberal employer, Mr. Henchman, I am indebted for complete specimens of its fructification, which enable me to establish it as a genus distinct from any yet described.
(*Footnote. Flinders Voyage volume 2 page 576.)
To this new genus I have given the name of my friend Captain King, who, during his important surveys of the Coasts of New Holland, formed valuable collections in several departments of Natural History, and on all occasions gave every assistance in his power to Mr. Cunningham, the indefatigable botanist who accompanied him. The name is also intended as a mark of respect to the memory of the late Captain Philip Gidley King, who, as Governor of New South Wales, materially forwarded the objects of Captain Flinders' voyage; and to whose friendship Mr. Ferdinand Bauer and myself were indebted for important assistance in our pursuits while we remained in that colony.
KINGIA.
ORD. NAT. Junceae prope Dasypogon, Calectasiam et Xerotem.
CHAR. GEN. Perianthium sexpartitum, regulare, glumaceum, persistens. Stamina sex, fera hypogyna: Antheris basi affixis. Ovarium triloculare, loculis monospermis; ovulis adscendentibus. Stylus 1. Stigma tridentatum. Pericarpium exsuccum, indehiscens, monospermum, perianthio scarioso cinctum.
Planta facie Xanthorrhoeae elatioris. Caudex arhorescens cicatricibus basibusve foliorum exasperatus? Folia caudicem terminantia confertissima longissima, figura et dispositione Xanthorrhoeae. Pedunculi numerosi foliis breviores, bracteis vaginantibus imbricatis tecti, floriferi terminales erecti, mox, caudice parum elongato foliisque novellis productis, laterales, et divaricati vel deflexi, terminati capitulo denso globoso floribus tribracteatis.
Kingia australis. Table C.
DESC. Caudex arborescens erectus simplicissimus cylindraceus, 6-18-pedes altus, crassitie femoris. Folia caudicem terminantia numerosissima patula, apicibus arcuato-recurvis, lorea, solida, ancipitia apice teretiusculo, novella undique tecta pilis adpressis strictis acutis laevibus, angulis lateralibus et ventrali retrorsum scabris. Pedunculi numerosi teretes 8-12-pollicares crassitie digiti, vaginis integris brevibus imbricatis hinc in foliolum subulatum productis tecti. Capitulum globosum, floridum magnitudine pruni minoris, fructiferum pomum parvum aequans. Flores undique dense imbricati, tribracteati, sessiles. Bractea exterior lanceolata breve acuminata planiuscula erecta, extus villosa intus glabra, post lapsum fructus persistens: duae laterales angusto-naviculares, acutissimae, carina lateribusque villosis, longitudine fere exterioris, simul cum perianthio fructifero, separatim tamen, dilabentibus. Perianthium sexpartitum regulare subaequale glumaceum: foliola lanceolata acutissima disco nervoso nervis immersis simplicissimis, antica et postica plana, lateralia complicata lateribus inaequalibus, omnia basi subangustata, extus longitudinaliter sed extra medium praecipue villosa, intus glaberrima, aestivatione imbricata. Stamina sex subaequalia, aestivatione stricta filamentis sensim elongantibus: Filamenta fere hypogyna ipsis basibus foliolorum perianthii quibus opposita leviter adhaerentia, filiformia glabra teretia: Antherae stantes, ante dehiscentiam lineares obtusae filamento paulo latiores, defloratae subulatae vix crassitie filamenti, loculis parallelo-contiguis connectivo dorsali angusto adnatis, axi ventrali longitudinaliter dehiscentibus, lobulis baseos brevibus acutis subadnatis: Pollen simplex breve ovale laeve. Pistillum: Ovarium sessile disco nullo squamulisve cinctum, lanceolatum trigono-anceps villosum, triloculare, loculis monospermis. Ovula erecta fundo anguli interioris loculi paulo supra basin suam inserta, obovata lenticulari-compressa, aptera: Testa in ipsa basi acutiuscula foramine minuto perforata: Membrana interna respectu testae inversa, hujusce nempe apici lata basi inserta, ovata apice angustato aperto foramen testae obturante: Nucleus cavitate membranae conformis, ejusdem basi insertus, caeterum liber, pulposus solidus, apice acutiusculo laevi aperturam membranae internae attingente. Stylus trigonus strictus, infra villosus, dimidio superiore glabro, altitudine staminum, iisdem paulo praecocior, exsertus nempe dum illa adhuc inclusa. Stigmata tria brevissima acuta denticuliformia. Pericarpium exsuccum, indehiscens, villosum, basi styli aristatum, perianthio scarioso et filamentis emarcidis cinctum, abortione monospermum. Semen turgidum obovatum retusum, integumento (testa) simplici membranaceo aqueo-pallido, bine (intus) fere a basi acutiuscula, raphe fusca verticem retusum attingente ibique in chalazam parvam concolorem ampliata. Albumen semini conforme dense carnosum album. Embryo monocotyledoneus, aqueo-pallidus subglobosus, extremitate inferiore (radiculari) acuta, in ipsa basi seminis situs, semi-immersus, nec albumine omnino inclusus.
Table C. figure 1. Kingiae australis pedunculus capitulo florido terminatus; figure 2, capitulum fructiferum; 3, sectio transversalis pedunculi: 4, folium: hae magnitudine naturali, sequentes omnes plus minus auctae sunt; 5, flos; 6, stamen; 7, anthera antice et, 8, eadem postice visa; 9, pistillum; 10, ovarii sectio transversalis; 11, ejusdem portio longitudinaliter secta exhibens ovulum adscendens cavitatem loculi replens; 12, ovulum ita longitudinaliter sectum ut membrana interna solummodo ejusque insertio in apice cavitatis testae visa sit; 13, ovuli sectio longitudinalis profundius ducta exhibens membranam internam et nucleum ex ejusdem basi ortum; 14, bracteae capituli fructiferi; 15, pericarpium perianthio filamentisque persistentibus cinctum; 16, pericarpium perianthio avulso filamentorum basibus relictis; 17, semen.
OBS. 1.
It remains to be ascertained, whether in this genus a resin is secreted by the bases of the lower leaves, as in Xanthorrhoea; and whether, which is probable, it agrees also in the internal structure of its stem with that genus. In Xanthorrhoea the direction of fibres or vessels of the caudex seems at first sight to resemble in some degree the dicotyledonous arrangement, but in reality much more nearly approaches to that of Dracaena draco, allowance being made for the greater number, and extreme narrowness of leaves, to which all the radiating vessels belong.*
(*Footnote. My knowledge of this remarkable structure of Xanthorrhoea is chiefly derived from specimens of the caudex of one of the larger species of the genus, brought from Port Jackson, and deposited in the collection at the Jardin du Roi of Paris by M. Gaudichaud, the very intelligent botanist who was attached to Captain De Freycinet's voyage.)
OBS. 2.
I have placed Kingia in the natural order Junceae along with Dasypogon, Calectasia and Xerotes, genera peculiar to New Holland, and of which the two former have hitherto been observed only, along with it, on the shores of King George's Sound.
The striking resemblance of Kingia, in caudex and leaves, to Xanthorrhoea, cannot fail to suggest its affinity to that genus also. Although this affinity is not confirmed by a minute comparison of the parts of fructification, a sufficient agreement is still manifest to strengthen the doubts formerly expressed of the importance of those characters, by which I attempted to define certain families of the great class Liliaceae.
In addition, however, to the difference in texture of the outer coat of the seed, and in those other points, on which I then chiefly depended in distinguishing Junceae from Asphodeleae, a more important character in Junceae exists in the position of the embryo, whose radicle points always to the base of the seed, the external umbilicus being placed in the axis of the inner or ventral surface, either immediately above the base as in Kingia, or towards the middle, as in Xerotes.
OBS. 3.
ON THE STRUCTURE OF THE UNIMPREGNATED OVULUM IN PHAENOGAMOUS PLANTS.
The description which I have given of the Ovulum of Kingia, though essentially different from the accounts hitherto published of that organ before fecundation, in reality agrees with its ordinary structure in Phaenogamous plants.
I shall endeavour to establish these two points; namely, the agreement of this description with the usual structure of the Ovulum, and its essential difference from the accounts of other observers, as briefly as possible at present; in tending hereafter to treat the subject at greater length, and also with other views.
I have formerly more than once* adverted to the structure of the Ovulum, chiefly as to the indications it affords, even before fecundation, of the place and direction of the future Embryo. These remarks, however, which were certainly very brief, seem entirely to have escaped the notice of those authors who have since written on the same subject.
(*Footnote. Flinders Voyage 2 page 601, and Linnean Society Transactions 12 page page 136.)
In the Botanical Appendix to the account of Captain Flinders' Voyage, published in 1814, the following description of the Ovulum of Cephalotus follicularis is given: Ovulum erectum, intra testam membranaceam continens sacculum pendulum, magnitudine cavitatis testae, and in reference to this description, I have in the same place remarked that, "from the structure of the Ovulum, even in the unimpregnated state, I entertain no doubt that the radicle of the Embryo points to the umbilicus."*
(*Footnote. Flinders Voyage loc. cit.)
My attention had been first directed to this subject in 1809, in consequence of the opinion I had then formed of the function of the Chalaza in seeds;* and sometime before the publication of the observation now quoted, I had ascertained that in Phaenogamous plants the unimpregnated Ovulum very generally consisted of two concentric membranes, or coats, enclosing a Nucleus of a pulpy cellular texture. I had observed also, that the inner coat had no connexion either with the outer or with the nucleus, except at its origin; and that with relation to the outer coat it was generally inverted, while it always agreed in direction with the nucleus. And, lastly, that at the apex of the nucleus the radicle of the future Embryo would constantly be found.
(*Footnote. Linnean Society Transactions 10 page 35.)
On these grounds my opinion respecting the Embryo of Cephalotus was formed. In describing the Ovulum in this genus, I employed, indeed, the less correct term sacculus, which, however, sufficiently expressed the appearance of the included body in the specimens examined, and served to denote my uncertainty in this case as to the presence of the inner membrane.
I was at that time also aware of the existence, in several plants, of a foramen in the coats of the Ovulum, always distinct from, and in some cases diametrically opposite to the external umbilicus, and which I had in no instance found cohering either directly with the parietes of the Ovarium, or with any process derived from them. But, as I was then unable to detect this foramen in many of the plants which I had examined, I did not attach sufficient importance to it; and in judging of the direction of the Embryo, entirely depended on ascertaining the apex of the nucleus, either directly by dissection, or indirectly from the vascular cord of the outer membrane: the termination of this cord affording a sure indication of the origin of the inner membrane, and consequently of the base of the nucleus, the position of whose apex is therefore readily determined.
In this state of my knowledge the subject was taken up in 1818, by my lamented friend the late Mr. Thomas Smith, who, eminently qualified for an investigation where minute accuracy and great experience in microscopical observation were necessary, succeeded in ascertaining the very general existence of the foramen in the membranes of the Ovulum. But as the foramina in these membranes invariably correspond both with each other and with the apex of the nucleus, a test of the direction of the future Embryo was consequently found nearly as universal, and more obvious than that which I had previously employed.
To determine in what degree this account of the vegetable Ovulum differs from those hitherto given, and in some measure, that its correctness may be judged of, I shall proceed to state the various observations that have been actually made, and the opinions that have been formed on the subject, as briefly as I am able, taking them in chronological order.
In 1672, Grew* describes in the outer coat of the seeds of many Leguminous plants a small foramen, placed opposite to the radicle of the Embryo, which, he adds, is "not a hole casually made, or by the breaking off of the stalk," but formed for purposes afterwards stated to be the aeration of the Embryo, and facilitating the passage of its radicle in germination. It appears that he did not consider this foramen in the testa as always present, the functions which he ascribes to it being performed in cases where it is not found, either, according to him, by the hilum itself, or in hard fruits, by an aperture in the stone or shell.
(*Footnote. Anatomy of Veget. begun page 3. Anatomy of Plants page 2.)
In another part of his work* he describes and figures, in the early state of the Ovulum, two coats, of which the outer is the testa; the other, his middle membrane, is evidently what I have termed nucleus, whose origin in the Ovulum of the Apricot he has distinctly represented and described.
(*Footnote. Anatomy of Plants page 210 table 80.)
Malpighi, in 1675,* gives the same account of the early state of the Ovulum; his secundinae externae being the testa, and his chorion the nucleus. He has not, however, distinguished, though he appears to have seen, the foramen of Grew, from the fenestra and fenestella, and these, to which he assigns the same functions, are merely his terms for the hilum.
(*Footnote. Anatome Plant. page 75 et 80.)
In 1694, Camerarius, in his admirable essay on the sexes of plants,* proposes, as queries merely, various modes in which either the entire grains of pollen, or their particles after bursting, may be supposed to reach and act upon the unimpregnated Ovula, which he had himself carefully observed. With his usual candour, however, he acknowledges his obligation on this subject to Malpighi, to whose more detailed account of them he refers.
(*Footnote. Rudolphi Jacobi Camerarii de sexu plantarum epistola page 8 46 et seq.)
Mr. Samuel Morland, in 1703,* in extending Leeuwenhoek's hypothesis of generation to plants, assumes the existence of an aperture in the Ovulum, through which it is impregnated. It appears, indeed, that he had not actually observed this aperture before fecundation, but inferred its existence generally and at that period, from having, as he says, "discovered in the seeds of beans, peas, and Phaseoli, just under one end of what we call the eye, a manifest perforation, which leads directly to the seminal plant," and by which he supposes the Embryo to have entered. This perforation is evidently the foramen discovered in the seeds of Leguminous plants by Grew, of whose observations respecting it he takes no notice, though he quotes him in another part of his subject.
(*Footnote. Philosophical Transactions volume 23 n. 287 page 1474.)
In 1704, Etienne Francois Geoffroy,* and in 1711, his brother Claude Joseph Geoffroy,** in support of the same hypothesis, state the general existence of an aperture in the unimpregnated vegetable Ovulum. It is not, however, probable that these authors had really seen this aperture in the early state of the Ovulum in any case, but rather that they had merely advanced from the observation of Grew, and the conjecture founded on it by Morland, whose hypothesis they adopt without acknowledgment, to the unqualified assertion of its existence, in all cases. For it is to be remarked, that they take no notice of what had previously been observed or asserted on the more important parts of their subject, while several passages are evidently copied, and the whole account of the original state and development of the Ovulum is literally translated from Camerarius' Essay. Nor does the younger Geoffroy mention the earlier publication of his brother, from which his own memoir is in great part manifestly derived.
(*Footnote. Quaestio Medica an Hominis primordia Vermis? in auctoris Tractatu de Materia Medica tome 1 page 123.)
(**Footnote. Mem. de l'Acad. des Sc. de Paris 1711 page 210.)
In 1718; Vaillant,* who rejects the vermicular hypothesis of generation, supposes the influence of the Pollen to consist in an aura, conveyed by the tracheae of the style to the ovula, which it enters, if I rightly understand him, by the funiculus umbilicalis: at the same time he seems to admit the existence of the aperture in the coat.
(*Footnote. Discours sur la Structure des Fleurs page 20.)
In 1745, Needham,* and in 1770, Gleichen,** adopt the hypothesis of Morland, somewhat modified, however, as they consider the particles in the grains of Pollen, not the grains themselves, to be the embryos, and that they enter the ovula by the umbilical cord.
(*Footnote. New Microscopical Discoveries page 60.)
(**Footnote. Observ. Microscop. page 45 et 61 paragraph 118.)
Adanson, in 1763,* states the Embryo to exist before fecundation, and that it receives its first excitement from a vapour or aura proceeding from the Pollen, conveyed to it through the tracheae of the style, and entering the Ovulum by the umbilical cord.
(*Footnote. Fam. des Plant. tom. 1 page 121.)
Spallanzani,* who appears to have carefully examined the unimpregnated Ovula of a considerable variety of plants, found it in general to be a homogeneous, spongy, or gelatinous body; but in two Cucurbitaceae to consist of a nucleus surrounded by three coats. Of these coats he rightly supposes the outermost to be merely the epidermis of the middle membrane or testa. Of the relative direction of the testa and inner coat in the two plants in question he takes no notice, nor does he in any case mention an aperture in the Ovulum.
(*Footnote. Fisica Anim. e Veget. tome 3 page 309 to 332.)
Gaertner, who, in the preface to his celebrated work, displays great erudition in every branch of his subject, can hardly, however, be considered an original observer in this part. He describes the unimpregnated Ovulum as a pulpy homogeneous globule, whose epidermis, then scarcely distinguishable, separates in a more advanced stage, and becomes the testa of the seed, the inner membrane of which is entirely the product of fecundation.* He asserts also that the Embryo constantly appears at that point of the ovulum where the ultimate branches of the umbilical vessels perforate the inner membrane; and therefore mistakes the apex for the base of the nucleus.
(*Footnote. Gaert. de Fruct. et Sem. 1 page 57, 59 et 61.)
In 1806 Mons. Turpin* published a memoir on the organ, by which the fecundating fluid is introduced into the vegetable ovulum. The substance of this memoir is, that in all Phaenogamous plants fecundation takes place through a cord or fasciculus of vessels entering the outer coat of the ovulum, at a point distinct from, but at the period of impregnation closely approximated to the umbilicus, and to the cicatrix of this cord, which itself is soon obliterated, he gives the name of Micropyle: that the ovulum has two coats, each having its proper umbilicus, or, as he terms it, omphalode; that these coats in general correspond in direction; that more rarely the inner membrane is, with relation to the outer, inverted; and that towards the origin of the inner membrane the radicle of the embryo uniformly points.
(*Footnote. Annal. du Mus. d'Hist. Nat. 7 page 199.)
It is singular that a botanist, so ingenious and experienced as M. Turpin, should, on this subject, instead of appealing in every case to the unimpregnated ovulum, have apparently contented himself with an examination of the ripe seed. Hence, however, he has formed an erroneous opinion of the nature and origin, and in some plants of the situation, of the micropyle itself, and hence also he has in all cases mistaken the apex for the base of the nucleus.
A minute examination of the early state of the ovulum does not seem to have entered into the plan of the late celebrated M. Richard, when in 1808 he published his valuable and original Analyse du Fruit. The ovulum has, according to him, but one covering, which in the ripe seed he calls episperm. He considers the centre of the hilum as the base, and the chalaza, where it exists, as the natural apex of the seed.
M. Mirbel, in 1815, though admitting the existence of the foramen or micropyle of the testa,* describes the ovulum as receiving by the hilum both nourishing and fecundating vessels,** and as consisting of a uniform parenchyma, in which the embryo appears at first a minute point, gradually converting more or less of the surrounding tissue into its own substance; the coats and albumen of the seed being formed of that portion which remains.***
(*Footnote. Elem. de Physiol. Veg. et de Bot. tome 1 page 49.)
(**Footnote. Id. tome 1 page 314.)
(***Footnote. Id. loc. cit.)
In the same year, M. Auguste de Saint Hilaire,* shows that the micropyle is not always approximated to the umbilicus; that in some plants it is situated at the opposite extremity of the ovulum, and that in all cases it corresponds with the radicle of the embryo. This excellent botanist, at the same time, adopts M. Turpin's opinion, that the micropyle is the cicatrix of a vascular cord, and even gives instances of its connexion with the parietes of the ovarium; mistaking, as I believe, contact, which in some plants unquestionably takes place, and in one family, namely, Plumbagineae, in a very remarkable manner, but only after a certain period, for original cohesion, or organic connexion, which I have not met with in any case.
(*Footnote. Mem. du Mus. d'Hist. Nat. 2 page 270 et seq.)
In 1815 also appeared the masterly dissertation of Professor Ludolf Christian Treviranus, on the development of the vegetable embryo,* in which he describes the ovulum before fecundation as having two coats: but of these, his inner coat is evidently the middle membrane of Grew, the chorion of Malpighi, or what I have termed nucleus.
(*Footnote. Entwick. des Embryo im Pflanzen-Ey.)
In 1822, Mons. Dutrochet, unacquainted, as it would seem, with the dissertation of Professor Treviranus, published his observations on the same subject.* In what regards the structure of the ovulum, he essentially agrees with that author, and has equally overlooked the inner membrane.
(*Footnote. Mem. du Mus. d'Hist. Nat. tome 8 page 241 et seq.)
It is remarkable that neither of these observers should have noticed the foramen in the testa. And as they do not even mention the well-known essays of MM. Turpin and Auguste de St. Hilaire on the micropyle, it may be presumed that they were not disposed to adopt the statements of these authors respecting it.
Professor Link, in his Philosophia Botanica, published in 1824, adopts the account given by Treviranus, of the coats of the ovulum before impregnation:* and of M. Turpin, as to the situation of the micropyle, and its being the cicatrix of a vascular cord. Yet he seems not to admit the function ascribed to it, and asserts that it is in many cases wanting.**
(*Footnote. Elem. Philos. Bot. page 338.)
(**Footnote. Id. page 340.)
The account which I have given of the structure of the vegetable ovulum, differs essentially from all those now quoted, and I am not acquainted with any other observations of importance respecting it.
Of the authors referred to, it may be remarked, that those who have most particularly attended to the ovulum externally, have not always examined it at a sufficiently early period, and have confined themselves to its surface: that those who have most minutely examined its internal structure, have trusted too much to sections merely, and have neglected its appearance externally: and that those who have not at all examined it in the early stage, have given the most correct account of its surface. This account was founded on a very limited observation of ripe seeds, generalized and extended to the unimpregnated ovulum, in connexion with an hypothesis then very commonly received: but this hypothesis being soon after abandoned, their statement respecting the ovulum was rejected along with it.
In the ovulum of Kingia, the inner membrane, with relation to the external umbilicus, is inverted; and this, as I have already observed, though in direct opposition to M. Turpin's account, is the usual structure of the organ. There are, however, several families in each of the two primary divisions of phaenogamous plants, in which the inner membrane, and consequently the nucleus, agrees in direction with the testa. In such cases the external umbilicus alone affords a certain indication of the position of the future embryo.
It is an obvious consequence of what has been already stated, that the radicle of the embryo can never point directly to the external umbilicus or hilum, though this is said to be generally the case by the most celebrated carpologists.
Another observation may be made, less obviously a consequence of the structure described, but equally at variance with many of the published accounts and figures of seeds, namely, that the radicle is never absolutely enclosed in the albumen; but, in the recent state, is either immediately in contact with the inner membrane of the seed, or this contact is established by means of a process generally very short, but sometimes of great length, and which indeed in all cases may be regarded as an elongation of its own substance. From this rule I have found one apparent deviation, but in a case altogether so peculiar, that it can hardly be considered as setting it aside.
It is necessary to observe, that I am acquainted with exceptions to the structure of the ovulum as I have here described it, In Compositae its coats seem to be imperforated, and hardly separable, either from each other or from the nucleus, in this family, therefore, the direction of the embryo can only be judged of from the vessels of the testa.* And in Lemna I have found an apparent inversion of the embryo with relation to the apex of the nucleus. In this genus, however, such other peculiarities of structure and economy exist, that, paradoxical as the assertion may seem, I consider the exception rather as confirming than lessening the importance of the character.
(*Footnote. Linnean Society Transactions 12 page 136.)
It may perhaps be unnecessary to remark, that the raphe, or vascular cord of the outer coat, almost universally belongs to that side of the ovulum which is next the placenta. But it is at least deserving of notice, that the very few apparent exceptions to this rule evidently tend to confirm it. The most remarkable of these exceptions occur in those species of Euonymus, which, contrary to the usual structure of the genus and family they belong to, have pendulous ovula; and, as I have long since noticed, in the perfect ovula only of Abelia.* In these, and in the other cases in which the raphe is on the outer side, or that most remote from the placenta, the ovula are in reality resupinate; an economy apparently essential to their development.
(*Footnote. Abel's China page 377.)
The distinct origins and different directions of the nourishing vessels and channel through which fecundation took place in the ovulum, may still be seen in many of those ripe seeds that are winged, and either present their margins to the placenta, as in Proteaceae, or have the plane of the wing at right angles to it, as in several Liliaceae. These organs are visible also in some of those seeds that have their testa produced at both ends beyond the inner membrane, as Nepenthes; a structure which proves the outer coat of scobiform seeds, as they are called, to be really testa, and not arillus, as it has often been termed.
The importance of distinguishing between the membranes of the unimpregnated ovulum and those of the ripe seed, must be sufficiently evident from what has been already stated. But this distinction has been necessarily neglected by two classes of observers. The first consisting of those, among whom are several of the most eminent carpologists, who have regarded the coats of the seed as products of fecundation. The second of those authors who, professing to give an account of the ovulum itself, have made their observations chiefly, or entirely, on the ripe seed, the coats of which they must consequently have supposed to be formed before impregnation.
The consideration of the arillus, which is of rare occurrence, is never complete, and whose development takes place chiefly after fecundation, might here, perhaps, be entirely omitted. It is, however, worthy of remark, that in the early stage of the ovulum, this envelope is in general hardly visible even in those cases where, as in Hibbertia volubilis, it attains the greatest size in the ripe seed; nor does it in any case, with which I am acquainted, cover the foramen of the testa until after fecundation.
The testa, or outer coat of the seed, is very generally formed by the outer membrane of the ovulum; and in most cases where the nucleus is inverted, which is the more usual structure, its origin may be satisfactorily determined; either by the hilum being more or less lateral, while the foramen is terminal; or more obviously, and with greater certainty where the raphe is visible, this vascular cord uniformly belonging to the outer membrane of the ovulum. The chalaza, properly so called, though merely the termination of the raphe, affords a less certain character, for in many plants it is hardly visible on the inner surface of the testa, but is intimately united with the areola of insertion of the inner membrane or of the nucleus, to one or other of which it then seems entirely to belong. In those cases where the testa agrees in direction with the nucleus, I am not acquainted with any character by which it can be absolutely distinguished from the inner membrane in the ripe seed; but as a few plants are already known, in which the outer membrane is originally incomplete, its entire absence, even before fecundation, is conceivable; and some possible cases of such a structure will be mentioned hereafter.
There are several cases known, some of which I have formerly noticed,* of the complete obliteration of the testa in the ripe seed; and on the other hand it appears to constitute the greater part of the substance of the bulb-like seeds of many Liliaceae, where it no doubt performs also the function of albumen, from which, however, it is readily distinguished by its vascularity.** But the most remarkable deviation from the usual structure and economy of the outer membrane of the ovulum, both in its earliest stage and in the ripe fruit, that I have yet met with, occurs in Banksia and Dryandra. In these two genera I have ascertained that the inner membrane of the ovulum, before fecundation, is entirely exposed, the outer membrane being even then open its whole length; and that the outer membranes of the two collateral ovula, which are originally distinct, cohere in a more advanced stage by their corresponding surfaces, and together constitute the anomalous dissepiment of the capsule; the inner membrane of the ovulum consequently forming the outer coat of the seed.
(*Footnote. Linnean Society Transactions 12 page 149.)
(**Footnote. Ibid.)
The inner membrane of the ovulum, however, in general appears to be of greater importance as connected with fecundation, than as affording protection to the nucleus at a more advanced period. For in many cases, before impregnation, its perforated apex projects beyond the aperture of the testa, and in some plants puts on the appearance of an obtuse, or even dilated stigma; while in the ripe seed it is often either entirely obliterated, or exists only as a thin film, which might readily be mistaken for the epidermis of a third membrane then frequently observable.
This third coat is formed by the proper membrane or cuticle of the Nucleus, from whose substance in the unimpregnated ovulum it is never, I believe, separable, and at that period is very rarely visible. In the ripe seed it is indistinguishable from the inner membrane only by its apex, which is never perforated, is generally acute and more deeply coloured, or even sphacelated.
The membrane of the nucleus usually constitutes the innermost coat of the seed. But in a few plants an additional coat, apparently originating in the inner membrane of Grew, the vesicula colliquamenti or amnios of Malpighi also exists.
In general the Amnios, after fecundation, gradually enlarges, till at length it displaces or absorbs the whole substance of the nucleus, containing in the ripe seed both the embryo and albumen, where the latter continues to exist. In such cases, however, its proper membrane is commonly obliterated, and its place supplied either by that of the nucleus, by the inner membrane of the ovulum, or, where both these are evanescent, by the testa itself.
In other cases the albumen is formed by a deposition of granular matter in the cells of the nucleus. In some of these cases the membrane of the amnios seems to be persistent, forming even in the ripe seed a proper coat for the embryo, the original attachment of whose radicle to the apex of this coat may also continue. This, at least, seems to me the most probable explanation of the structure of true Nymphaeaceae, namely, Nuphar, Nymphaea, Euryale, Hydropeltis, and Cabomba, notwithstanding their very remarkable germination, as observed and figured in Nymphaea and Nuphar by Tittmann.*
(*Footnote. Keimung der Pflanzen page 19 et 27 table 3 et 4.)
In support of this explanation, which differs from all those yet given, I may here advert to an observation published many years ago, though it seems to have escaped every author who has since written on the subject, namely, that before the maturity of the seed in Nymphaeaceae, the sacculus contains along with the embryo a (pulpy or semi-fluid) substance, which I then called Vitellus, applying at that time this name to every body interposed between the albumen and embryo.* The opinion receives some confirmation also from the existence of an extremely fine filament, hitherto overlooked, which, originating from the centre of the lower surface of the sacculus, and passing through the hollow axis of the Albumen, probably connects this coat of the Embryo in an early stage with the base of the nucleus.
(*Footnote. Prodr. Flor. Nov. Holl. 1 page 306.)
The same explanation of structure applies to the seeds of Piperaceae and Saururus; and other instances occur of the persistence either of the membrane or of the substance of the amnios in the ripe seed.
It may be concluded from the whole account which I have given of the structure of the ovulum, that the more important changes consequent to real, or even to spurious fecundation, must take place within the nucleus: and that the albumen, properly so called, may be formed either by a deposition or secretion of granular matter in the utriculi of the amnios, or in those of the nucleus itself, or lastly, that two substances having these distinct origins, and very different textures, may co-exist in the ripe seed, as is probably the case in Scitamineae.
On the subject of the ovulum, as contained in an ovarium, I shall at present make but one other remark, which forms a necessary introduction to the observations that follow.
ON THE STRUCTURE OF THE FEMALE FLOWER IN CYCADEAE AND CONIFERAE.
That the apex of the nucleus is the point of the ovulum where impregnation takes place, is at least highly probable, both from the constancy in the appearance of the embryo at that point, and from the very general inversion of the nucleus; for by this inversion its apex is brought nearly, or absolutely, into contact with that part of the parietes of the ovarium, by which the influence of the pollen may be supposed to be communicated. In several of those families of plants, however, in which the nucleus is not inverted, and the placentae are polyspermous, as Cistineae,* it is difficult to comprehend in what manner this influence can reach its apex externally, except on the supposition, not hastily to be admitted, of an impregnating aura filling the cavity of the ovarium; or by the complete separation of the fecundating tubes from the placentae, which, however, in such cases I have never been able to detect.
(*Footnote. This structure of ovulum, indicated by that of the seed, as characterizing and defining the limits of Cistineae (namely, Cistus, Helianthemum, Hudsonia and Lechea) I communicated to Dr. Hooker, by whom it is noticed in his Flora Scotica (page 284) published in 1821; where, however, an observation is added respecting Gaertner's description of Cistus and Helianthemum, for which I am not accountable.)
It would entirely remove the doubts that may exist respecting the point of impregnation, if cases could be produced where the ovarium was either altogether wanting, or so imperfectly formed, that the ovulum itself became directly exposed to the action of the pollen, or its fovilla; its apex, as well as the orifice of its immediate covering, being modified and developed to adapt them to this economy.
But such, I believe, is the real explanation of the structure of Cycadeae, of Coniferae, of Ephedra, and even of Gnetum, of which Thoa of Aublet is a species.
To this view the most formidable objection would be removed, were it admitted, in conformity with the preceding observations, that the apex of the nucleus, or supposed point of impregnation, has no organic connexion with the parietes of the ovarium. In support of it, also, as far as regards the direct action of the pollen on the ovulum, numerous instances of analogous economy in the animal kingdom may be adduced.
The similarity of the female flower in Cycadeae and Coniferae to the ovulum of other phaenogamous plants, as I have described it, is indeed sufficiently obvious to render the opinion here advanced not altogether improbable. But the proof of its correctness must chiefly rest on a resemblance, in every essential point, being established, between the inner body in the supposed female flower in these tribes, and the nucleus of the ovulum in ordinary structures; not only in the early stage, but also in the whole series of changes consequent to fecundation. Now as far as I have yet examined, there is nearly a complete agreement in all these respects. I am not entirely satisfied, however, with the observations I have hitherto been able to make on a subject naturally difficult, and to which I have not till lately attended with my present view.
The facts most likely to be produced as arguments against this view of the structure of Coniferae, are the unequal and apparently secreting surface of the apex of the supposed nucleus in most cases; its occasional projection beyond the orifice of the outer coat; its cohesion with that coat by a considerable portion of its surface, and the not unfrequent division of the orifice of the coat. Yet most of these peculiarities of structure might perhaps be adduced in support of the opinion advanced, being apparent adaptations to the supposed economy.
There is one fact that will hardly be brought forward as an objection, and which yet seems to me to present a difficulty, to this opinion; namely, the greater simplicity in Cycadeae, and in the principal part of Coniferae, of the supposed ovulum which consists of a nucleus and one coat only, compared with the organ as generally existing when enclosed in an ovarium. The want of uniformity in this respect may even be stated as another difficulty, for in some genera of Coniferae the ovulum appears to be complete.
In Ephedra, indeed, where the nucleus is provided with two envelopes, the outer may, perhaps, be supposed rather analogous to the calyx, or involucrum of the male flower, than as belonging to the ovulum; but in Gnetum, where three envelopes exist, two of these may, with great probability, be regarded as coats of the nucleus; while in Podocarpus and Dacrydium, the outer cupula, as I formerly termed it,* may also, perhaps, be viewed as the testa of the ovulum. To this view, as far as relates to Dacrydium, the longitudinal fissure of the outer coat in the early stage, and its state in the ripe fruit, in which it forms only a partial covering, may be objected.** But these objections are, in a great measure, removed by the analogous structure already described in Banksia and Dryandra.
(*Footnote. Flinders Voyage volume 2 page 573.)
(**Footnote. Id. loc. cit.)
The plurality of embryos sometimes occurring in Coniferae, and which, in Cycadeae, seems even to be the natural structure, may also, perhaps, be supposed to form an objection to the present opinion, though to me it appears rather an argument in its favour.
Upon the whole, the objections to which the view here taken of the structure of these two families is still liable, seem to me, as far as I am aware of them, much less important than those that may be brought against the other opinions that have been advanced, and still divide botanists on this subject.
According to the earliest of these opinions, the female flower of Cycadeae and Coniferae is a monospermous pistillum, having no proper floral envelope.
To this structure, however, Pinus itself was long considered by many botanists as presenting an exception.
Linnaeus has expressed himself so obscurely in the natural character which he has given of this genus, that I find it difficult to determine what his opinion of its structure really was. I am inclined, however, to believe it to have been much nearer the truth than is generally supposed; judging of it from a comparison of his essential with his artificial generic character, and from an observation recorded in his Praelectiones, published by Giseke.*
(*Footnote. Praelect. in Ord. Nat. page 589.)
But the first clear account that I have met with, of the real structure of Pinus, as far as regards the direction, or base and apex of the female flowers, is given, in 1767, by Trew, who describes them in the following manner: "Singula semina vel potius germina stigmati tanquam organo feminino gaudent,"* and his figure of the female flower of the Larch, in which the stigmata project beyond the base of the scale, removes all doubt respecting his meaning.
(*Footnote. Nov. Act. Acad. Nat. Curios. 3 page 453 table 13 figure 23.)
In 1789, M. de Jussieu, in the character of his genus Abies,* gives a similar account of structure, though somewhat less clearly as well as less decidedly expressed. In the observations that follow, he suggests, as not improbable, a very different view, founded on the supposed analogy with Araucaria, whose structure was then misunderstood; namely, that the inner scale of the female amentum is a bilocular ovarium, of which the outer scale is the style. But this, according to Sir James Smith,** was also Linnaeus' opinion; and it is the view adopted in Mr. Lambert's splendid monograph of the genus published in 1803.
(*Footnote. Gen. Pl. page 414.)
(**Footnote. Rees Cyclop. art. Pinus.)
In the same year in which Mr. Lambert's work appeared, Schkuhr* describes, and very distinctly figures, the female flower of Pinus, exactly as it was understood by Trew, whose opinion was probably unknown to him.
(*Footnote. Botan. Handb. 3 page 276 table 308.)
In 1807, a memoir on this subject, by Mr. Salisbury, was published,* in which an account of structure is given, in no important particular different from that of Trew and Schkuhr, with whose observations he appears to have been unacquainted.
(*Footnote. Linnean Society Transactions 8 page 308.)
M. Mirbel, in 1809,* held the same opinion, both with respect to Pinus and to the whole natural family. But in 1812, in conjunction with M. Schoubert,** he proposed a very different view of the structure of Cycadeae and Coniferae, stating, that in their female flowers there is not only a minute cohering perianthium present, but an external additional envelope, to which he has given the name of cupula.
(*Footnote. Ann. du Mus. d'Hist. Nat. tome 15 page 473.)
(**Footnote. Nouv. Bulletin des Sc. tome 3 pages 73, 85 et 121.)
In 1814 I adopted this view, as far, at least, as regards the manner of impregnation, and stated some facts in support of it.* But on reconsidering the subject, in connexion with what I had ascertained respecting the vegetable ovulum, I soon after altogether abandoned this opinion, without, however, venturing explicitly to state that now advanced, and which had then suggested itself.**
(*Footnote. Flinders Voyage 2 572.)
(**Footnote. Tuckey Congo page 454 et Linnean Society Transactions volume 13 page 213.)
It is well known that the late M. Richard had prepared a very valuable memoir on these two families of plants; and he appears, from some observations lately published by his son, M. Achille Richard,* to have formed an opinion respecting their structure somewhat different from that of M. Mirbel, whose cupula is, according to him, the perianthium, more or less cohering with the included pistillum. He was probably led to this view, on ascertaining, which I had also done, that the common account of the structure of Ephedra was incorrect,** its supposed style being in reality the elongated tubular apex of a membranous envelope, and the included body being evidently analogous to that in other genera of Coniferae. |
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