The carpology and taxonomic relationships of Bretschneidera (Bretschneideraceae)
DOI:
https://doi.org/10.24310/abm.v21i0.8671Keywords:
Carpology, Bretschneidera, Bretschneideraceae, phylogeny, seedAbstract
The fruit and seed anatomy and morphology of Bretschneidera sinensis Hemsl., the monotypic genus in the Bretschneideraceae, have been studied in order to clarify its phylogenetic relationships. The loculicidal trilocular syncarpous fruit has unusual unspecialised, aerenchymatous fibrous 12-14-layered endocarp and sclerified innermost part of the mesocarp with numerous radially elongated bundles pervading the parenchyma of mesocarp. Such a distribution of mechanical elements is indicative of the derivation of the Bretschneidera fruit from a drupaceous fruit of the «durian» type. In order to emphasise hard differences in pericarpic mechanical system of between drupe, capsule and this specialised type it is proposed to establish a new type of fruits - ascade (aseada, lat.) - to accommodate all fruits originated from the superior ovary with sclerified mechanical system located in the mesocarp. Thus the fruit of Bretschneidera is defined as a loculicidal ascade instead of loculicidal capsule. Seeds of Bretschneidera are relatively large, practically exalbuminous, with straight dicotyledonous embryo. The spermoderm of Bretschneidera is exomesotestal with unspecialised crushed tegmen, 40-45-layered, traversed by 8 postchalazal vascular bundles. Evidence mainly from seed morphology and anatomy of seed coats emphasises the somewhat anomaly of the traditional inclusion of Bretschneideraceae in the Sapindales, being quite distinct in spermoderm structure and origin from both Sapindaceae and Hippocastanaceae as well as from other families of the order, excepting anomalous exo-mesotestal Akaniaceae. Furthermore, seed anatomy does not confirm any relationships with Capparales. It is suggested that Bretschneideraceae with Akaniaceae constitute a distinct relict side-branch of connaraceous-sapindaceous ancestry. It can by no means be considered as a basal clade to Capparales. It is suggested to keep Bretschneideraceae with Akaniaceae separately from Sapindales and Capparales before more comprehensive studies of seedcoat anatomy of other orders of Rosidae.
Downloads
Metrics
References
BOESEWINKEL, F.D. -1990- Ovule and seed development of Tovaria pendula Ruiz et Pavon. Bot. Jahrb. Syst., I ll: 389-401.
BOUFFORD, D. E., KJAER, A., MADSEN, J. O. & SKRYDSTRUP, T. -1989- Glucosinolates in Bretschneideraceae. Biochem Syst. Ecol., 17: 375-379.
CHAW, S.-M. & PENG, C.-I. -1987- Palynological notes on Bretschneidera sinensis Hemsl. Bol. Bull. Acad. Sin., 28: 55-60.
CORNER, E. J. H. -1976- The seeds ofdicotyledons. Vols. 2. Cambridge: Cambridge University Press.
CRONQUIST, A. -1992- An integrated system of classification of flowering plants. 2nd ed. New York: Columbia University Press.
CZERNIAKOWSKAYA, E. F. -1992.- Resedaceae. Pp. 202-208 in A. L. Takhtaj an (Ed.), Anatomia Seminum Comparativa. Vol. 4. St.-Petersburg: Nauka. [In Russian].
DAHLGREN, G. -1989- The last Dahlgrenogram. System of classification of the dicotyledons. Pp. 249-260 in K. Tan (Ed.), Plant taxonomy, phytogeography and related subjects. The Davis and Hedge Festschrift. Edinburgh: Edinburgh University Press.
DAHLGREN, R. -1983- General aspects of angiosperm evolution and macrosystemat cs. Nord. J. Bol., 3: 119-149.
DOWELD, A. B. -1996a- Staphyleaceae, Tapisciaceae, Sapindaceae, Hippocastanaceae, Melianthaceae, Akaniaceae. Pp. 306-341, 343-349 in A. L. Takhtajan (Ed.), Anatomia Seminum Comparativa. Vol. 5. St.-Petersburg: Nauka. [In Russian].
DOWELD, A. B. -1996b- Zaklyucheniye k poriadku Sapindales [Conclusions on the order Sapindales]. Pp. 349-350 in A. L. Takhtajan (Ed.), Anatomia Seminum Comparativa. Vol. 5. St.-Petersburg: Nauka [In Russian].
DOWELD, A. B. -1996c- The systematic relevance of fruit and seed anatomy and morphology of Akania (Akaniaceae). Bot. J. Linn. Soc., 120: 379-389.
ENGLER, A. -1919- Syllabus der Pflanzenfamilien; cine Übersicht iiber das gesamte Pflanzensystem mit besonderer Berticksichtigung der Medizinalund Nutzpflanzen, nebst einer Übersicht u?ber die Florenreiche und Florengebiete der Erde, zum gebrauch bei Vorlesungen und Studien u?ber spezielle und medizinisch-pharmazeutische Botanik. 8 Aufl. Berlin: Gebrtider Borntraeger.
ENGLER, A. - i 924- Syllabus der Pflanzenfamilien; cine Obersicht u?ber das gesamte Pflanzensystem mit besonderer Beriicksichtigung der Medizinalund Nutzpflanzen, nebst einer Ubersicht tiber die Florenreiche und Florengebiete der Erde, zum gebrauch bei Vorlesungen und Studien tiber spezielle und medizinisch-pharmazeutische Botanik. 9 & 10 Aufl. Berlin: Gebriider Born traeger.
GADEK, P.A., E.S. FERNANDO, C.J. QUINN, S.B. HOOT, T. TERRAZAS, M.C. SHEANAN & M.W. CHASE -1996- Sapindales: molecular delimitation and infraordinal groups. Amer. J. Bot., 83: 802-811.
GADEK, P. A., C.J. QUINN„ J.E. RODMAN, K.G. KAROL, E. CONTI, R.A. PRICE, & F.S. FERNANDO -1992- Affinities of the australian endemic Akaniaceae: new evidence from the rbeL sequences. Austr. Syst. Bot., 15: 717-724.
GARCIN, A.-G. - 1891 - Contribution a l'étude des péricarpes charnus. Du noyau des drupes. Histologic et histogénese. Ann. Soc. Bot. Lyon, 17 (1890): 27-39.
GOLDBERG, A. -1986- Classification, evolution, and phylogeny of the families of dicotyledons. Smiths. Contr. Bot., 58: 1-314.
GUERIN, P. -1901- Développement de la graine et en particulier du tegument seminal de quelques Sapindacées. J. Bot. (Morot), 15: 336-362.
HARBOURNE, J.B., B.L. TURNER -1984- Plant chemosystematics. London: Academic Press.
HEIMSCH, C. -1942- Comparative anatomy of the secondary xylem in the «Gruinales» and «Terebinthales» of Wettstein with reference to taxonomic grouping. Lilloa, 8: 83-198.
HEMSLEY, W.B. -1901- Bretschneidera sinensis. Hooker's Icon. Pl. Ser. 4. 8(28): 2708.
HUTCHINSON, J. -1926- The families of flowering plants. I. Dicotyledons. Arranged according to a new system based on their probable phylogeny. London: MacMillan & Co.
KADEN, N.N. -1964- Osnovy evolutsionnoy morfologii plodov [Fundamentals of the evolutionary morphology of fruits]. Vol. 1. Doct. Sci. Diss. Moscow: Moscow M. V. Lomonosov State University, Dept. of Higher Plants. [In Russian].
KOMAR, G.A., G.B. RODIONOVA -1992- Moringaceae. Pp. 209-212 in A. L. Takhtajan (Ed.), Anatomía Seminum Comparativa. Vol. 4. St.-Petersburg: Nauka. [In Russian].
LE MONNIER, G. -1872- Recherches sur la nervation de la graine. Ann. Sci. Nat. Bot. Sér. 5, 16: 233-305.
LECLERC DU SABLON, M. -1884- Sur la déhiscence du fruits á péricarpe sec. Ann. Sci. Nat. Bot. Sér. 6, 18: 5-104.
LIU, C. -1986- Studies of pollen morphology in the Bretschneideraceae and the relative families. Acta Bot. Yunnan., 8: 441-450. [In Chinese].
LÜ, J. & Y.S. HU -1994- Structural study of secondary xylem in Bretschneidera sinensis Hemsl. Acta Bot. Sin., 36: 459-465. [In Chinese].
LÜ, J. & Y.-X. HU -1994- Anatomy of the young stem and secondary phloem in Bretschneidera sinensis Hemsl. Chin. J. Bot., 6: 112-117.
MAGEN, K. -1912- Beitrdge zur vergleichenden Anatomic der Samenschalen einiger Familien aus der Engler'schen Reihe der Sapindales. Inaug.-Diss. Univ. Zu?rich. Borna-Leipzig: Buchdruckerei R. Noske.
O'BRIEN, T.P. & M.E. MCCULLY -1981- The study of plant structure: principles and selected methods. Melbourne: Termarcarphi and Pty. Ltd.
RADLKOFER, L. -1908- Bretschneidera. Pp. 208- 209 in A. Engler & K. Prantl (Eds), Die natu?rlichen Pflanzenfamilien. Nachtrage 3. Leipzig: W. Englemann.
RODIONOVA, G.B. -1992a- Capparaceae. Pp. 183- 191 in A. L. Takhtajan (Ed.), Anatómia Seminum Comparativa. Vol. 4. St.-Petersburg: Nauka. [In Russian].
RODIONOVA, G.B. -1992b- Tovariaceae. P. 202 in A. L. Takhtajan (Ed.), Anatomia Seminum Comparativa. Vol. 4. St.-Petersburg: Nauka. [In Russian].
RODIONOVA, G.B. -1992c- Brassicaceae. Pp. 191- 201 in A. L. Takhtajan (Ed.), Anatomia Seminum Comparativa. Vol. 4. St.-Petersburg: Nauka. [In Russian].
RODMAN, J.E. -1991a- A taxonomic analysis of glucosinolate-producing plants. Part I. Phenetics. Syst. Bol., 16: 589-618.
RODMAN, J.E. -1991b- A taxonomic analysis of glucosinolate-producing plants. Part 2. Cladistics. Syst. Bot., 16: 619-629.
RODMAN J.E., R.A. PRICE, K. KAROL, E. CONTI, K.J. SYTSMA & J.D. PALMER -1993- Nucleotide sequences of the rbcL gene indicate monophyly of mustard oil plants. Ann. Missouri Bot. Gard., 80: 686-699.
SCHMID, R. -1986- On Cornerian and other terminology of angiospermous and gymnospermous seed coats: historical perspective and terminological recommendations. Taxon, 35: 476-491.
SPJUT, R.W. -1994- A systematic treatment of fruit types. Mem. N.Y. Bol. Gard., 70: 1-182.
TAKHTAJAN, A.L. -1987- Systema Magnoliophytorum. Leningrad: Nauka. [In Russian].
TANG, W. -1935- Timber studies of Chinese trees. VII. Notes on the systematic position of Bretschneideraceae as shown by its timber anatomy. Bull. Fan Mem. Inst. Biol., 6: 153-159.
THORNE, R.F. -1992- An updated phylogenctic classification of the flowering plants. Aliso, 13: 365-389.
TOBE, H. & C.-I. PENG -1990- The embryology and taxonomic relationships of Bretschneidera (Bretschneideraceae). Bot. J. Linn. Soc., 103: 139-152.
TOBE, H. & P.H. RAVEN -1995- Embryology and relationships of Akania (Akaniaceae). Bot. J. Linn. Soc., 118: 261-274.
YANG, D.-Q. &C.-M. HU -1985- The chromosomes of Bretschneidera Hemsl. Notes Roy. Bot. Gard. Edinb., 42: 347-349.
WEBERBAUER, A. -1898- Beitrdge zur Anatomie der Kapselfru?chte. Bol. Centralbl., 73: 54-59, 81-105, 135-142, 161-168, 193-202, 250-257, 296-302.
ZENG, M.X. & Z.L. TONG -1984- Anatomical study of the stem of Bretschneidera sinensis. S.W. Agrie. Coll. Bull., 1984(1): 42-46. [In Chinese].
Downloads
Published
How to Cite
Issue
Section
License
Those authors who publish in this journal accept the following terms:
a. The authors will retain their copyrights and guarantee the journal the right of first publication of their work, which will be simultaneously subject to the Creative Commons Attribution-Non-commercial 4.0 license whose full text can be found at <http: // creative commons .org / licenses / by-nc / 4.0> that allows third parties to share the work as long as its author and its first publication are indicated, and as long as it is not for commercial purposes.
b. Authors may adopt other non-exclusive licensing agreements for the distribution of the version of the published paper (e.g., deposit it in an institutional telematic file or publish it in a monographic volume) provided that the initial publication in this journal be indicated.
c. Authors are allowed and recommended to disseminate their work through the Internet (e.g., in institutional telematic archives or on their websites) before and during the submission process, which can produce interesting exchanges and increase citations of the published work. (See The effect of open access)