Flora Australiensis/Volume 5/Proteaceae

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Order CIV. PROTEACEAE

Flowers hermaphrodite or rarely partially unisexual. Perianth regular or irregular, deciduous, consisting of 4 segments valvately united in the bud, the claws forming a tube cylindrical or dilated towards the base, the laminæ short, forming a globular ovoid or rarely elongated limb; the segments at length separating either from the base upwards or revolute from the laminæ downwards, leaving a portion of the tube entire or open on one side, the laminæ sometimes cohering long after the segments have separated lower down. Stamens 4, opposite the perianth-segments and usually inserted on them, either with the filaments wholly adnate leaving the anthers sessile at the base of the laminæ, or the filaments shortly free below the laminæ, or very rarely the stamens, entirely free from the perianth; anthers various, all perfect or rarely partially abortive, most frequently with 2 parallel cells adnate to a connectivum continuous with the filament. Hypogynous or perigynous glands or scales in many genera 4, alternating with the stamens, but in some genera variously united or reduced in number or wholly deficient. Ovary 1-celled, sessile or stipitate, more or less excentrical, with a single terminal undivided style, variously shaped at the end, with a small terminal oblique or lateral stigma. Ovules either solitary, or 2 collaterally attached or slightly superposed, or several imbricate in 2 contiguous rows, either pendulous and orthotropous, or more frequently laterally attached and more or less amphitropous, rarely erect and anatropous, the micropyle always inferior and frequently prominent from the incomplete development of the primine. Fruit either an indehiscent nut or drupe, or a more or less dehiscent coriaceous or woody follicle, very rarely a completely 2-valved capsule; either 1-celled and 1-seeded, or when 2 seeds are ripened in a drupe sometimes really 2-celled from the growth of the endocarp between as well as round the seeds, or when 2 or more seeds ripen in a follicle, apparently 2- or more-celled by the consolidation of the external coating of the 2 adjoining seeds into a membranous or woody plate detaching itself from the remainder of the seed. Seeds without albumen, the testa usually thin, rarely coriaceous or hard; embryo straight, with fleshy cotyledons and a short inferior radicle.—Shrubs or trees, rarely undershrubs or even perennial herbs. Leaves alternate or scattered, in a very few genera strictly, opposite or verticillate, but often crowded under the inflorescence so as to appear verticillate, usually coriaceous, often vertical with stomata on both sides, or in the same genera horizontal or narrow and terete, entire toothed or variously divided, without stipules. Flowers axillary or terminal, solitary or in racemes or spikes, often condensed into umbels heads or cones, each flower or pair of flowers subtended by a bract, very deciduous in some genera and perhaps sometimes really deficient, the pedicels always without bracteoles.

Proteaceæ, with their chief seat in Australia and South Africa, extend on the one hand to New Caledonia, the Indian Archipelago, and tropical Asia, chiefly eastern, to Japan, and on the other to South America. The seven tribes of the Order are all in Australia. Of tha first four, constituting the Nucamentaceæ, the two principal ones, Proteeæ and Personiæ, are also in South Africa, but represented by different genera, the nine Australian ones being, as well as the four constituting the small tribes Conospermeæ and Franklandieæ, all endemic with the exception of a single New Zealand species of Persoonia, and a New Caledonian Canarrhenes. None of the Nucamentaceæ are either in America or Asia, for the South American Andripetalum and Guevina, referred by Meissner to Persoonieæ, belong with Helicia and Macadamia to the Grevilleæ. Of the Australian genera of this tribe of Grevilleeæ, Helicia is chiefly Asiatic, Adenostephanus is tropical American, with one New Caledonian species, and the large genus Grevillea has also a few New Caledonian species, the remaining eight genera are endemic. Of the Australian Embothrieæ, Lomatia extends to the Andes of South America, where it is accompanied by two nearly allied genera, and Stenocarpus to New Caledonia; the two remaining genera, as well as the two which constitute the tribe Banksieæ, are endemic in Australia.

The clavate fusiform or disk-shaped end of the style in Proteaceæ is usually described as the stigma, and when it is more or less constricted it is said to be articulate, but I have never found any real articulation, and although the thickened style-end may be an essential aid in the collection or dissemination of the pollen, its surface is not stigmatic, the real stigma being usually very small, either on the point terminating the style-end; or in the centre of the disk, or quite lateral. The diversified mode in which in different genera the conformation of this style-end and its relation to the anthers promotes the dissemination of pollen whilst it impedes self-fertilization, upon which I have drawn up a few notes for the Linnean Society founded on the examination of dried specimens, would be an interesting study for local botanists who have the means of examining and watching the plants living in their native stations.

In the distribution of the numerous species of this most natural Order into tribes, genera, and sections, I have only had to follow, with slight modifications, the admirable arrangement proposed by Brown and further developed by Endlicher and Meissner; but in the great subdivision into Nucamentaceæ and Folliculares, these terms most not be taken strictly in their literal sense, for indehiscent drupes occur in both divisions. Taking however the fruit generally, in conjunction with the arrangement of the ovules and the inflorescence, neither of them again strictly constant, we have very fairly definite characters for two large groups which are both natural and to a certain degree geographical. For although both are abundant in Australia, the Nucamentaceæ alone are in Africa, and the Folliculares alone in Asia and America.

Suborder 1. Nucamentaceæ.Fruit an indehiscent nut or drupe. Flowers usually solitary within each bract.

Tribe 1. Proteeæ.Anthers all perfect or very rarely the upper one abortive, with 2 parallel cells adnate to the connectivum, inserted at the base of the short spreading laminæ of the perianth. Ovule 1 or rarely 2. Stigma terminal. Fruit a dry nut.

Flowers in dense cone-like spikes or beads with imbricate scale-like bracts, with few or many outer empty bracts firming an involucre. Anthers free.

Cone-scales firmly adhering to the rhachis and opening for the emission of the more or less flattened nuts.

1. Petrophila

Cone-scales either very deciduous or remaining closely imbricate after flowering till they fall off with the nuts which are not flattened.

2. Isopogon

Flowers solitary within an involucre of 4 to 8 bracts

3. Adenanthos

Flowers in small heads with very small bracts. Anthers cohering round the style and the adjoining cells of two different anthers applied face to face in the bud forming a single cell.

4. Stirlingia

Tribe 2. Conospermeæ.Anthers: one with 2 perfect cells, two with 1 perfect and 1 abortive cell, the fourth abortive, the perfect cells broad, concave, erect, without any connective, the adjoining ones of distinct anthers applied face to face in the bud forming a single cell; all on very short thick filaments at the base of the laminæ or at the summit of the tube of the perianth. Ovule 1. Fruit a dry nut.

Upper anthers abortive, replaced by a short membrane connecting the filament with the disk-shaped stigma. Nut ovoid or oblong. Leaves mostly divided.

5. Synaphea

Lower anther abortive. Stigma raised above the stamens on the beak-like end of the style. Nut turbinate, flat and comose on the top. Leaves entire.

6. Conospermum

Tribe 3. Franklandieæ.Anthers all perfect with parallel adnate cells enclosed in and adnate to the slender perianth-tube. Ovule 1. Fruit a dry nut with a poppus-like cone.

Single genus

7. Franklandia

Tribe 4. Persoonieæ.Anthers all perfect with parallel cells adnate to the connective, the stamens inserted at or below the middle of the perianth-segments. Ovules 2 or sometimes 1. Fruit a drupe or rarely a dry nut or membranous.

Leaves divided or lobed. Flowers in interrupted spikes or racemes. Ovules 2. Fruit dry, indehiscent.

Filaments inserted on the perianth, converging end united in a ring round the style. Fruit a nut.

8. Symphyonema

Stamens free at the base of the perianth-segments. Fruit membranous, flattened

9. Bellendena

Leaves entire. Flowers in interruptsd axillary spikes. Ovule 1. Fruit a 3-winged nut.

10. Agastachys

Leaves toothed. Flowers in interrupted axillary spikes. Ovule 1. Fruit a drupe.

11. Cenarrhenes

Leaves entire. Flowers axillary or rarely forming a terminal or infra-terminal raceme by the abortion of the floral leaves. Ovules 2 or 1. Fruit a drupe.

12. Persoonia

Suborder 2. Folliculares.Fruit dehiscent, follicular or 2-valved, rarely (in the first 2 genera), drupaceous and indehiscent. Flowers usually in pairs, with a single bract to each pair, rarely (in Carnarvonia, Lambertia and Stenocarpus, the inflorescence anomalous.

Tribe 5. Grevilleeæ.Ovules 2 or 4, collateral. Seeds without any intervening substances or separated by a thin lamina or meaty substance. Flowers in racemes or clusters, with deciduous or abortive bract; or with an involucre of imbricate bracts.

Ovules 2. Perianth regular or nearly so, small (under ½ in. except in one species), the anthers on short filaments attached below the laminæ. Style cylindrical or clavate at the end.

Flowers pedicellate, in pairs, in racemes. Fruit with a thick woody indehiscent pericarp or putamen.

Leaves alternate. Ovules ascending.

13. Helicea

Leaves verticillate. Ovules descending

14. Macadamia

Flowers sessile, in pairs, in cylindrical spikes. Fruit thick and woody, tardily dehiscent. Leaves opposite

15. Xylomelum

Flowers solitary or clustered on irregularly branched peduncles. Fruit a follicle. Leaves alternate, compound.

16. Carnarvonia

Flowers sessile or nearly so, in pairs, in cylindrical or oblong spikes. Fruit a follicle. Leaves alternate.

17. Orites

Ovules 2. Perianrh long and narrow. Anthers linear. Flowers solitary, or 7 together in an involucre of persistent imbricate bracts. Leaves verticillate.

18. Lambertia

Ovules 2. Perianth revolute in the bud or rarely straight and regular. Anthers short and sessile within the concave laminæ. Leaves alternate.

Ovules orthotropous, pendulous. (Fruit a drupe?).

19. Adenostephanus

Ovules amphitropous, laterally attached. Fruit a follicle.

Seeds without wings or the wings short at both ends or annular. Inflorescence terminal, rarely also axillary.

20. Grevillea

Seeds winged, chiefly or entirely at the upper end. Inflorescence axillary.

21. Hakea

Ovules 4, collateral. Perianth revolute in the bud or straight and regular. Anthers short and sessile within the concave lamin&aelig. Fruit a follicle. Leaves alternate.

Perianth revolute in the bud. Hypogynous gland unilateral or semiannular. Follicle short and broad.

22. Buckinghamia

Perianth straight. Hypogynous glands 4. Follicle oblong, recurved

23. Darlingia
Tribe 6. Embothrieæ.Ovules several, imbricate in 2 rows. Seeds usually separated by thin laminæ or a mealy substance.

Flowers in short compact racemes, surrounded by an involucre of imbricate coloured bracts.

24. Telopea.

Flowers in loose racemes. Bracts small or deciduous.

Hypogynous glands 3. Ovules imbricate upwards. Seeds winged at the upper end.

25. Lomatia.

Hypogynous glands 4. Ovules imbricate downwards. Seeds with narrow wings all round

26. Cardwellia.

Flowers in umbels without bracts.

27. Stenocarpus.
Tribe 7. Banksieæ.Ovules 2, collateral. Seeds separated either by a hard usually woody substance or by a membrane rarely wanting. Flowers in dense cones or heads.

Flowers in ovoid or cylindrical cones, without any involucre.

28. Banksia

Flowers in heads surrounded by an involucre of imbricate bracts and floral leaves.

29. Dryandra

Suborder 1. Nucamentaceæ—Fruit an indehiscent nut or drupe, either 1-seeded or if 2-seeded the seeds separated by a complete dissepiment continuous with the endocarp. Flowers usually solitary within each bract, in cones or spikes or solitary, very rarely racemose, the spikes often shortened into heads.

Tribe 1. Proteeæ—Anthers all perfect, or very rarely the upper one abortive, with 2 parallel cells adnate to the connectivum, inserted at the base of the short spreading lamin&aelig of the perianth. Ovule 1 or in a very few species a second one more or less developed. Stigma at the point of the straight style end. Fruit a dry nut.

Petrophila
Isopogon
Adenanthos
Stirlingia


Tribe 2. Conospermeæ.—Anthers: one with 2 perfect cells, two with 1 perfect and 1 abortive cell, the fourth abortive, the perfect cells broad, concave, erect, without any connective, the adjoining ones of distinct anthers applied face to face, so as to form in the bud one cell; all on very short thick filaments at the base of the laminæ or summit of the perianth-tube. Ovule 1. Fruit a dry nut.

Synaphea
Conospermum


TRIBE 3. FRANKLANDIEÆ.—Anthers all perfect with adnate parallel cells, enclosed in and adnate to the slender perianth-tube. Ovule 1. Fruit a dry nut with a pappus-like coma.

Franklandia


TRIBE 4. PERSOONIEÆ.—Anthers all perfect, with parallel cells adnate to the connective, the stamens inserted at or below the middle of the perianth-segments. Ovules 2, or sometimes 1. Fruit a drupe or rarely a dry nut or membranous.

Symphonema
Bellendena
Agastachys
Cenarrhenes
Persoonia


SUBORDER 2. FOLLICULARES.—Fruit dehiscent, follicular or 2-valved or rarely drupaceous and indehiscent. Flowers usually in pairs, with a single bract to each pair, or rarely the inflorescence anomalous. Ovules 2 or more, collateral in each pair.

TRIBE 5. GREVILLEEÆ.—Ovules 2 or 4, all collateral. Seeds without any intervening substances or separated only by a thin lamina or mealy substance. Flowers in racemes or rarely in umbels or clusters, with deciduous or abortive bracts or rarely surrounded by an involucre or imbricate bracts.

Helicia
Macadamia
Xylomelum
Carnarvonia
Orites
Lambertia
Adenostephanus
Grevillea
Hakea
Buckinghamia
Darlingia


TRIBE 6. EMBOTHRIEÆ.—Ovules several, imbricate in 2 rows. Seeds usually separated by thin laminæ or a mealy substance, (possibly the outer coating of the seeds detached and united as in Banksia).

Telopea
Lomatia
Cardwellia
Stenocarpus


TRIBE 7. BANKSIEÆ.—Ovules 2, collateral. Seeds separated by a hard or membranous, usually bifid, sometimes double plate, rarely wanting. Flowers in dense spikes or cones with closely imbricate persistent bracts within or below the spike.

The singular plate intervening between the two seeds in this tribe has been explained by Brown to consist of the outer coating of one side of each seed, separating from the inner coatings as they advance towards maturity, the two becoming usually consolidated opposite the nuclei, remaining distinct opposite the seed-wings. This plate is entirely free from the walls of the pericarp, except at the point of attachment of the seed, forming a portion of the latter, not of the former, and has therefore no title to the name of a dissepiment, real or spurious, still given to it in systematic works, evin in the Prodromus.

Banksia
Dryandra