Hornschuchia (Annonaceae), an endemic and threatened genus from the Brazilian Atlantic Forest Author Vilela, Lucas Universidade de São Paulo, Instituto de Biociências, Departamento de Botânica, Rua do Matão, 277, ed. Sobre-as-Ondas, 05508 - 090 São Paulo, SP, Brazil. & Universidade Anhembi-Morumbi, Escola de Ciências da Saúde, Rua Dr. Almeida Lima, 1134, 03101 - 001, Mooca, São Paulo, SP, Brazil. vilela.lucas@hotmail.com Author Lopes, Jenifer De Carvalho Universidade de São Paulo, Instituto de Biociências, Departamento de Botânica, Rua do Matão, 277, ed. Sobre-as-Ondas, 05508 - 090 São Paulo, SP, Brazil. jenifer.clopes@gmail.com text European Journal of Taxonomy 2022 2022-07-11 828 75 108 http://dx.doi.org/10.5852/ejt.2022.828.1859 journal article 10.5852/ejt.2022.828.1859 c5427427-07fc-4449-ba86-25f19cd47482 2118-9773 6823766 Hornschuchia bryotrophe Nees Figs 1–2 Flora 4 (1): 302 ( Nees 1821 ) . Type : BRAZIL Bahia • “Ilhéus”, “ Strasse des Capitains Filisberto ” [via Felisbertia ]; Dec. 1816 ; M . A . P . zu Wied s.n. ( 34, Nees von Esenbeck nr. 34 ); holotype : BR [ BR0000006593025 ]!; isotypes: BR [ BR0000006593353 ]!, M [ M0240081 ]!, MEL [ MEL2123975 ]!, NY [ NY00008358 ] !. Heterotypic synonym Mosenodendron insigne R.E.Fr., Kongliga Svenska Vetenskaps Akademiens Handlingar , ser. 3 34 (5): 8 ( Fries 1900 ) . Type : BRAZIL Bahia • “ Vittoria ”; F. Sellow s.n. ; holotype : B [ B 10 0243155 ]!. Selected material examined BRAZIL Bahia Gandú , Estrada a Itamari ”; [ 13°43′24.9″ S , 39°35′13.3″ W ]; 22 Dec. 1970 ; T . S . Santos 1168 ; CEPEC [ CEPEC00006203 ]!, NY [ NY00395782 ] ! • Guaratinga , ca 2.5 Km na estrada Itabela-Guaratinga , saindo da BR-101. Entrada em estrada de terra à direita, no sentido Itabela-Guaratinga ”; 16°33′48.2″ S , 39°45′09.5″ W ; alt. 145 m ; 24 Jan. 2009 ; M . Groppo et al. 1808 ; SPF ! • Ibicaraí , Rodovia BR-415, 40 km Oeste de Itabuna. Região de Mata Higrográfila Sul Baiana ”; [ 14°51′27.7″ S , 39°31′09.8″ W ]; alt. 300 m ; S . A . Mori s.n. et al. ; RB [ RB00042146 ] ! • Ilhéus ”; Jan. 1837 ; B . Luschnath s.n. ; BR [ BR0000006592691 ] ! • “Castelnovo” [ Castelo Novo ]; [ 14°38′34.6″ S , 39°12′28.0″ W ]; Riedel 711 ; M [ M0240082 ] ! • Ipiaú , Rodovia Ipiaú-Ibirataia ”; [ 14°06′21.4″ S , 39°40′39.8″ W ]; 13 Nov. 1971 ; T . S . Santos 2133 ; CEPEC [ CEPEC00008059 ]!, NY [ NY00395783 ] ! • Itamaraju , Assentamento Pedra Bonita , aproximadamente 20 Km da rodovia vicinal de Itamaraju sentido Jucuruçu ”; 16°50′19″ S , 39°37′53″ W ; alt. 500 m ; 13 Feb. 2007 ; R . A . X. Borges 769 ; CEPEC [ CEPEC00116346 ] ! • Maraú , Rod. BR 030 , trecho Ubaitaba / Marau , km 15. Vegetação perturbada em Região de Mata Hidrográfica Sul baiana. Folha SD 24 (14-39c)”; [ 14°15′29.0″ S , 39°12′28.0″ W ]; S . A . Mori s.n. et al. ; RB [ RB00042161 ] ! • Una , Reserva Biológica de Una ; REBIO de Una , Entrada no km 46da rodovia BA001 Ilhéus-Una ”; [ 15°06′34.5″S , 39°00′11.5″ W ]; 23 Jan. 2006 ; J . L . Paixão 661 et al. ; RB [ RB00495855 ] ! • São Felipe , Recôncavo Sul , Serra da Capioba ”; 12°47′05.7″ S , 39°04′06.7″ W ; 1956; R . P . Lôrdelo 56-333 ; ALCB [ ALCB004360 ]!. – Espírito Santo Águia Branca , Rochedo , Trilha do Córrego , proprietário Ailton Corteleti ”; 18º57′21″ S , 40º48′5″ W ; alt. 300–400 m ; 19 Dec. 2007 ; V . Demuner et al. 4816 ; MBML [ MBML032572 ] ! • Águas Claras , Escola Agroecológica ”; 18º53′32″ S , 40º43′48″ W ; alt. 300–500 m ; 1 Feb. 2006 ; L . F . S . Magnago et al. 531 ; MBML [ MBML027865 ]!, SPF ! • Aracruz , Aldeia Candeias ”; [ 19°46′38.6″ S , 40°12′02.8″ W ]; 26 Mar. 1997 ; M . A . de Assis et al. 903 ; SPF ! • Comboios ”; [ 19°44′40.1″ S , 40°00′00.7″ W ]; 27 Jul. 1992 ; O . J . Pereira 3692 ; VIES [ VIES008830 ] ! • Picuã ”; [ 19°50′31.4″ S 40°19′58.0″ W ]; 4 Jun. 2011 ; C . L . Dalmonech et al. 50 ; MBML [ MBML044614 ] ! • Cariacica , Reserva Biológica Duas Bocas Localidade de Duas Bocas , Trilha do Pescador ”; 20°16′44″ S , 40°28′42″ W ; alt. 135 m ; 20 Oct. 2008 ; P . H . Labiak et al. 5000 ; CEPEC [ CEPEC00129318 ]!, MBM [ MBML037745 ]!, RB [ RB00544259 ] ! • Guarapari , estrada ES-477, que liga a BR-101 à rodovia do Sol (ES-060), Ca de 6 km da BR 101 , Mata ao lado da estrada, entrada pela Fazenda Bonanza ”; 20°31′48″ S , 40°25′12″ W ; alt. 30–50 m ; 17 Feb. 1999 ; R . Mello-Silva et al. 1596 ; CEPEC [ CEPEC00083584 ]!, MBM [ MBM 235550 ]!, NY [ NY01145439 ]!, NY [ NY01145438 ]!, RB [ RB00042129 ]!, RB [ RB00042117 ]!, SPF ! • “Linhares” (Jaguaré*), “ Barra Seca , Fazenda São Carlos , propriedade do Sr. Aleixo Barnabé , ca 1.4 km E da BR 101 ”; [ 18°59′09.8″ S , 39°59′24.4″ W ]; 20 Feb. 1995 ; J . R . Pirani et al. 3579 ; RB [ RB00042105 ]!, SPF !, VIES [ VIES037620 ] ! • Reserva Natural da Companhia Vale do Rio Doce ”; 19.0069° S , 40.1661° W ; 2 Mar. 2010 ; A . Q . Lobão 1532 ; SPF !, SAMES [ SAMES00216 ]!, VIES [ VIES024740 ] ! • Marilândia , Estrada não pavimentada para São Rafael. Mata de encosta, ao lado direito da estrada”; 19°24′42.5″ S , 40°28′34.2″ W ; alt. 200 m ; 19 Jan. 2011 ; P . Fiaschi et al. 3478 ; MBML [ MBML042352 ]!, RB [ RB00681608 ]!, SPF ! • Montanha , Fazenda Luis Siqueira - distrito da Penha”; 18°8′45.14″ S , 40°19′1.8″ W ; alt. 140 m ; 15 Nov. 2012 ; A . M . Assis et al. 3515 ; MBML [ MBML047032 ]! • ( Governador Lindenberg *) Fazenda São Jorge , 18 km from church in Bananal on paved road from Bananal to Novo Brazil , behind house; [ 19°14′29.8″ S , 40°26′41.7″ W ]; 21 Apr 1995 ; J . A . Kallunki et al. 707 ; NY [ NY00227611 ]!, SPF ! • Santa Leopoldina , Bragança , Rancho Chapadão , proprietário: João Emilio ”; 20°7′22.8″ S , 40°32′46.8″ W ; alt. 280–550 m ; 30 Mar. 2006 ; V . Demuner et al. 2101 ; MBML [ MBML026395 ]!, SPF ! • Sooretama , Reserva Biológica de Sooretama. Porção Oeste , trilha do Barro Roxo. Floresta de Tabuleiro ”; [ 19°01′29.2″ S , 40°08′05.4″ W ]; 18 Jan. 2010 ; A . Q . Lobão et al. 1542 ; SAMES [ SAMES00217 ] ! • Vitória , Junção BR-101/ES-060, Rod. para Praia do Sol , 6 Km BR-101, 3 Km ES-060”; [ 20°31′19.0″ S , 40°23′25.2″ W ]; 31 Jul. 1991 ; D. M . Johnson et al. 1847 ; CEPEC [ CEPEC00064494 ]!, SPF !. – Minas Gerais Serra do Cipó. Serra do Cipó ”; [ 19°20′32.6″ S 43°35′54.1″ W ]; 16 Jan. 1951 ; J . G Kuhlmann et al. s.n. ; RB [ RB00042164 ]!. – Pernambuco Ipojuca , Engenho de Conceição Velha , Mata das Três Passagens , à direita da PE-60, em direção a Barreiras , após a tubulação que cruza a estrada para a Usina Ipojuca ”; [ 8°23′31.7″ S , 35°05′26.3″ W ]; 9 Dec. 1997 ; S . Tavares et al. 51 ; NY [ NY01017923 ]!. – Rio de Janeiro Rio de Janeiro ”; J . G . Kuhlmann s.n. ; RB [ RB00042140 ] ! • Taunay , Morro de mata baixa”; [ 22°57′31.6″ S , 43°16′37.8″ W ]; 27 Feb. 1951 ; G . A . Black et al. 51-11753 ; IAN [ IAN069581 ] ! • “Estrada do Tanguá, perto de Rio Bonito. Rio Bonito” ( Cachoeiras de Macacu *); [ 22°39′37.5″ S , 42°45′10.7″ W ]; 26 Feb. 1951 ; J . G Kuhlmann et al. s.n. ; RB [ RB00042134 ]!, RB [ RB 150900]!, US [ US 01346584] ! • Cultivada no Jardim Botânico do Rio de Janeiro”; RB [ RB00042137 ]!. – Locality unknown C.F.P. von Martius , Herb. Fl. Bras. n. 858 ; M [ M0240084 ] !. Additional material examined BRAZIL V. Demuner 3220 ; MBML [ MBML09241 ]!, SPF ! V. Demuner 4412 ; MBML [ MBML032022 ]!, SPF ! J.H.L. El Ottra 131 ; SPF ! D.A. Folli 1699 ; SPF ! 2917 ; RB [ RB00484637 ]!, SPF ! C.N. Fraga 2480 ; CEPEC [ CEPEC00130062 ]!, MBML [ MBML038885 ]!, RB [ RB00557920 ]! F.S. Gomes 1539 ; ALCB [ ALCB049674 ]! E. Guimarães 182 ; RB [ RB00042094 ]! J.G. Jardim 262 ; NY [ NY00395787 ]! J.G. Jardim 623 ; CEPEC [ CEPEC00064214 ]!; J.G. Jardim 8852 ; RB [ RB00852603 ]!, SPF ! J.A. Kallunki 589 ; K [ K001191170 ]!, NY [ NY00395780 ]!, SPF ! J.A. Kallunki 696 ; NY [ NY00227526 ]!, SPF ! J.G. Kuhlmann s.n. ; NY [ NY00395784 ]! J.C. Lopes 111 ; SPF ! J.C. Lopes 151 ; SPF ! P.J.M. Maas 8829 ; MBML [ MBML009585 ]!, NY [ NY02699059 ]!, RB [ RB00484639 ]! P.J.M. Maas 9811 ; NY [ NY02699058 ]!, SPF ! L.F.S. Magnago 489 ; MBML [ MBML030986 ]! L.F.S. Magnago 1646 ; MBML [ MBML027722 ]!, SPF ! R. Mello-Silva 1174 ; NY [ NY NY00395781 ]!, NY [ NY00395777 ]!, RB [ RB00042117 ]!, SPF ! R. Mello-Silva 3141 ; SPF ! R.F. Monteiro 329 ; RB [ RB00516860 ]!, SPF ! S.A. Mori 9292 ; CEPEC [ CEPEC00013256 ]! S.A. Mori 9368 ; NY [ NY00395785 ]! S.A. Mori 11348 ; CEPEC [ CEPEC00015681 ]!, NY [ NY00395786 ]! O.J. Pereira 4019 ; VIES [ VIES008836 ]! O.J. Pereira 2472 ; VIES [ VIES006348 ]! J.R. Pirani 3074 ; K [ K001191168 ]!, K [ K001191167 ]!, MBM [ MBM181229 ]!, NY [ NY00395778 ]!, SPF ! Without collectorʼs information ; RB [ RB00042137 ]! C. Rocini 7 ; SPF ! L.A.M. Silva 1555 ; CEPEC [ CEPEC00029042 ]! W.W. Thomas 3720 ; CEPEC [ CEPEC00090940 ]!, NY [ NY00684336 ]! W.W. Thomas 10742 ; CEPEC [ CEPEC00064423 ]!, MBM [ MBM187065 ]!, NY [ NY00095141 ]! W.W. Thomas 11069 ; CEPEC [ CEPEC00069957 ] ! • W.W. Thomas 12654 ; CEPEC [ CEPEC00091555 ]!. Description Shrubs or trees, 0.5–5 m tall. Leaves chartaceous; petiole 0.5–7 mm long; lamina 5.6–35 × 1.9–9.5 cm , narrowly oblong to elliptic, narrowly obovate to oblanceolate, both surfaces glabrous, base asymmetric, acute to obtuse, apex acuminate to acute, obtuse or rarely emarginate; primary vein impressed adaxially and raised abaxially, 6–22 pairs of secondary veins, commissural veins evident on both sides, impressed adaxially and raised abaxially, angles between primary and secondary veins 50–60°. Inflorescence 6–22-flowered; flagelliflorous, inflorescence branched into axes 12–60 cm long, running near the soil, rarely trunciflorous, ramiflorous or terminal inflorescence; bracts persistent, 5.5–9(–13) × (1–)1.5–3(–3.5) mm. Flowers with pedicel 1–40 mm long; flower buds 1–12 × 1–3 mm , conical to ellipsoid, densely covered in trichomes. Sepals completely connate, calyx cupuliform, apex truncate, 1–4 × 1–6 mm , glabrescent. Petals linear, white, rarely pinkish, 11–19 mm long, densely covered in trichomes at the base and glabrous to glabrescent towards the apex; stamens 6, 3–6 × 0.5 mm ; carpels 3, 4–9 × 0.5 mm . Monocarps 1–3, fusiform, 1.3–7 × 0.1–0.9 cm , glabrous, green in vivo; stipe 3–5 mm long, calyx caducous. Seeds 2–6, oblong-ellipsoid, 11–20 × 3.5–6 mm , brownish, rugose, with aril. Distribution and habitat Hornschuchia bryotrophe has the widest distribution, occurring from the States of Pernambuco , Bahia , Espírito Santo to Rio de Janeiro ( Fig. 1 ). The species has been collected only once in Pernambuco State , close to the coast in the Atlantic Forest ( Bazante & Alves 2021 , Tavares et al. 51 ; NY[NY01017923]). There is a single collection that has been made in Serra do Cipó, Cerrado biome (Mello-Silva et al. 2012). However, this is the only record in the whole genus outside the Atlantic Forest and it is possible that this was a mistake in the label annotation (Mello-Silva et al. 2012, Kuhlmann et al. s.n. ; RB[RB00042164]!). Hornschuchia bryotrophe occurs mainly in moist forest, but also in seasonal semideciduous forest ( Fig. 1 ). In Bahia , it inhabits lowland tropical moist forest ( Gouvêa et al. 1976 ; Thomas & Barbosa 2008 ) that occurs near the coast up to 70 km inland ( Mori & Silva 1979 ). In Espírito Santo , it occurs in the northern portion of the state, in tabuleiro (ʻtablelandʼ) forest, a seasonal semideciduous forest ( Peixoto et al. 2008 ). Phenology Flowering from January to December and fruiting from December to July. Conservation status Least Concern (BGCI & IUCN 2019). Notes There are two type collections of Hornschuchia bryotrophe at BR, for one of them, the holotype , Nees von Esenbeck annotated the number 34 ([BR0000006593025], Moraes et al. 2016 ), the same annotation is also on the isotypes at the M and MEL herbaria. The type localities of both H. bryotrophe and H. myrtillus , “Strasse des Capitains Filisberto”, known as via Felisbertia, was a road linking Ilhéus with Minas Gerais whose construction was ordered and paid for by Marechal Felisberto Caldeira Brant ( Moraes et al. 2016 ). The work of Gallagher & Moraes (2014) explains how the specimens collected by Wied in Brazil , such as the type collections of H. bryotrophe and H. myrtillus , arrived in an Australian herbarium, MEL. Hornschuchia bryotrophe is one of the species with cauliflory. The development of cauliflory begins in the axil of the leaf. However, it remains active even after the abscission of the spent inflorescence branch due to the presence of dormant buds that repeatedly produce new inflorescences ( Endress 2010 ). Cauliflory is classified into different types : (i) ramiflory refers to the inflorescence emerging in leafless portions of the branch; (ii) trunciflory to inflorescences borne on the main trunk; (iii) basiflory to inflorescences produced at the base of the trunk that could either be sessile or on perennial brachyblasts ( Mildbraed 1922 ). In basiflory, the flower axis may elongate, which is called idiocladanthy ( Mildbraed 1922 ). In cases in which the flower-bearing axis in idiocladanthy further elongates, allowing the inflorescence branches to reach the soil and grow along the ground, the term flagelliflory is applied ( Mildbraed 1922 ). In such cases, the inflorescence emerges from the base of the trunk in long compound branches close to the soil. In H. bryotrophe , three types of cauliflory have been observed: flagelliflory, ramiflory and trunciflory. However, the term flagelliflory is also adopted for flowers or inflorescences that are exposed from the canopy of the tree in long rope-like branches associated with bat pollination (Weberling 1989: 233–234). Fig. 2. Morphological characters of Hornschuchia bryotrophe Nees. A . Habit , showing the cauliflorous inflorescence emerging from the base of the trunk with inflorescence branches highly ramified ( J.C. Lopes 111 , SPF). B . Inflorescence branch with floral buds ( J.R. Pirani 3074 , SPF). C . Flower (L.F.S. Magnago 1646 , SPF). D . Monocarp ( C. Roccini 7 , SPF). Drawing by Laura Montserrat. Hornschuchia bryotrophe is distinct from the other species of the genus by its leaves with an evident commissural vein ( Fig. 2B ). Hornschuchia bryotrophe , H. polyantha and H. myrtillus present flagelliflory ( Fig. 2A ). Nevertheless, in addition to the leaves, H. bryotrophe also differs from these species by the conical flower buds with acute apex ( Fig. 2B ) (vs cylindrical with obtuse apex).