Phytotaxa 637 (3): 251–260 ISSN 1179-3155 (print edition)

https://www.mapress.com/pt/
Article PHYTOTAXA
Copyright © 2024 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.637.3.3

A new species of Compsoneura (Myristicaceae) from interandean valleys of middle

Magdalena and lower Cauca rivers, Colombia
Boris Villanueva Tamayo1,3 & Álvaro Cogollo Pacheco2,4
1
Jardín Botánico de Bogotá “José Celestino Mutis”, Av José Celestino Mutis #68-95, Bogotá, Colombia.
� bvillanueva@jbb.gov.co; � bsvillanuevat@ut.edu.co; https://orcid.org/0000-0002-6929-3572
2
Jardín Botánico Joaquín Antonio Uribe de Medellín, Calle 73 51D-14, Medellín, Colombia.
� cogolloi@yahoo.com; https://orcid.org/0000-0002-3885-0815
3
Universidad del Tolima, Grupo de Investigación de Biodiversidad y Dinámica de Ecosistemas Boscosos-GIBDET.
4
Corporación Salvamontes Colombia, Medellín Colombia.

Abstract

Compsoneura choibo, a new species of Myristicaceae from lowland wet forests located in interandean valleys in middle
Magdalena (inhabited by Ateles hybridus) and lower Cauca rivers, Colombia, is described, illustrated, and its morphological
relationships with allied species is discussed. This new species is similar to C. mutisii, but it is characterized by the fruits
with smooth pericarp but densely covered by T-shaped trichomes. A morphological description, taxonomic comments, an
illustration, information about its current conservation status and an updated key to species of Compsoneura are presented.
Compsoneura has its center of distribution in Colombia, where 12 of the species are known to occur.

Key words: Endemic species, lowlands, Ateles, tree

Resumen

Compsoneura choibo, una nueva especie de Myristicaceae de bosques húmedos de tierras bajas de los valles interandinos del
medio río Madgalena (con presencia de Ateles hybridus) y bajo río Cauca, Colombia, es descrita, ilustrada y se discuten sus
relaciones morfológicas con las especies aliadas. Esta nueva especie es similar a C. mutisii, pero se caracteriza por el pericarpo
liso pero cubierto con tricomas en forma de T. Se presenta una descripción morfológica, comentarios taxonómicos, una
ilustración, información acerca de su estado actual de conservación y se incluye una clave de las especies de Compsoneura.
De estas, 12 se conocen en Colombia, el país con la mayor diversidad del género.

Palabrasclave: árbol, especies endémicas, Ateles, tierras bajas

Introduction

The Myristicaceae Brown (1810: 399) includes 21 genera (Santamaria & Lagomarsino 2022). It is mostly distributed in
humid lowland forests of the world’s tropics (Kühn & Kubitzki 1993). The family is primarily composed of dioecious
species, with the only exception of Iryanthera (A.DC.) Warburg (1897: 126) in the Americas, which includes some
monoecious species (Smith & Wodehouse 1937). These plants are typically trees or shrubs, although occasionally they
can be found as vines, as in the case of Pycnanthus Warburg (1897: 252).
Identifying members of this family vegetatively is relatively straightforward due to a combination of distinctive
features. These features include a hyaline exudate ranging in color from red to dark red, orange, and amber. Additionally,
they have simple, alternate leaves and a characteristic whorled and branching growth pattern, which is known as the
“Massart model” of Myristicaceae tree architecture (Hallé et al. 1978).
The neotropical genus Compsoneura (A.DC.) Warburg (1897: 125) stands out among other genera of the family
due to its distinctive characteristics, such as the leaves with tertiary veins that are conspicuously subparallel, somewhat
perpendicular to both the midrib and secondary veins. What further sets Compsoneura apart is the nature of its pericarps,
which can take two forms: they may be very thin and notably dehiscent, or remarkably woody, thick, often sulcate, and

Accepted by Pedro Fiaschi: 6 Feb. 2024; published: 22 Feb. 2024 251

sometimes carinate, or unusually smooth (Warburg 1897, Gentry 1993, Janovec 2000, Rodrigues et al. 2001, Aymard
et al. 2020).
Compsoneura was initially proposed as a section of Myristica Gronovius (1755: 141) by De Candolle (1856),
with three species: C. capitellata (A.DC.) Warburg (1897: 146), C. debilis (Spruce ex A.DC.) Warburg (1897: 144),
and C. sprucei (A.DC.) Warburg (1897: 143). Otto Warburg, in his Monograph of Myristicaceae (1897), categorized
Compsoneura in two sections: C. section Eucompsoneura Warburg (1897: 143) comprising species with free anthers,
including C. sprucei, C. debilis, and C. capitellata, and C. section Coniostele Warburg (1897: 147), featuring species
with fused anthers, such as C. trianae Warburg (1897: 147).
Smith and Wodehouse (1937) recognized eight species of Compsoneura following the two sections proposed by
Warburg (1897), and synonymized C. costaricensis Warburg (1905: 71) under C. sprucei, and C. tessmannii Markgraf
(1926: 964) under C. capitellata. Later, Smith (1950) published C. cuatrecasasii A.C.Smith (1950: 318) and a few
years later, Virola atopa Smith (1950: 328) was transferred to C. atopa (A.C.Sm.) A.C.Smith (1956: 573). Other
important contributions were made by Rodrigues (1989), with the publication of two species: C. rigidifolia Rodrigues
(1989: 160) and C. schultesiana Rodrigues (1989: 161), and Jaramillo & Baslev (2001), who proposed C. lapidiflora
Jaramillo & Balslev (2001: 561), a new species from Ecuador. Janovec (2002, 2003), increased our knowledge of
the genus by publishing three new species: C. diazii Janovec (2002: 366), C. anoriensis Janovec & A.K.Neill (2003:
254) and C. claroensis Janovec & A.K.Neill (2003: 257). He also proposed C. mexicana (Hemsl.) Janovec (2002:
668) as a valid species for Mexico and Central America floras. Lately, C. nallarettiana M.Ríos, R.Zárate & J.Grandez
(2023: 294) was described from Peru. Currently, the genus comprises 18 accepted species (including the new species
described here).
The present contribution increases to 12 the number of Compsoneura species known from Colombia, which is
the center of distribution of this genus, where 67% of the species are known to occur. This new species was identified
during research in the lowland humid forests of the middle Magdalena and lower Cauca river basins in Colombia.

Materials and methods

The specimens of Compsoneura collected were confronted with specimens from the following herbaria: BOG, COAH,
COL, CUVC, FMB, HPUJ, HUA, JBB, MEDEL, TOLI, UDBC (acronyms follow Thiers 2019, continuously updated).
In addition, specialized literature was consulted (Warburg 1897, Warburg 1905, Smith & Wodehouse 1937, Smith
1938, Smith 1950, Smith 1956, Janovec 2000, Jaramillo & Baslev 2001, Janovec & Harrison 2002, Janovec & Neill
2003, Aymard et al. 2020; Ríos et al. 2023), as well as the images of nomenclatural types available at Global Plants
on JSTOR (https://plants.jstor.org/). The illustration was done by freehand, and the photographs of male flowers were
taken from samples preserved in ethanol 50%, used a Nikon D7000 camera with a 105 mm macro lens and a Raynox
250 magnifying glass. The morphological measurements were carried out using a discovery digital caliper. A Lankester
Composite Digital Plate (LCDP) was prepared. To determine the conservation status according to IUCN categories
and criteria (IUCN 2017), the extent of occurrence (EOO) and area of occupancy (AOO) were calculated using the
Geospatial Conservation Assessment Tool (GeoCat) (Bachman et al. 2011), constantly updated through the https://
geocat.kew.org/

Taxonomy

Compsoneura choibo Villanueva & Cogollo, sp. nov. (Figs. 1–2)

Type:—COLOMBIA. Santander: Municipality of Cimitarra, Hacienda Bocas del San Juan, Caño San Juan, 6°41’48’’N, 74°7’12.9’’W,
115 m, 19 April 2016 (fl ♂), B. Villanueva-T., J. Cabezas & V. Beltran 1897 (holotype: JBB! [accession 39801]; isotypes: ANDES!,
COL!, JAUM!, JBB!, TOLI!, UDBC).
Diagnosis:—This new species can be distinguished from other species of Compsoneura by the eucamptodromous venation of the leaves,
the tomentose fruits, non-carinate fruits, and seeds with a very short red aril. It is somewhat similarity to C. mutisii, but this the new
species differs by its tomentose fruits with ferruginous trichomes, a rostrate apex, and seeds with a reduced red aril at the base.

252 • Phytotaxa 637 (3) © 2024 Magnolia Press VILLANUEVA-TAMAYO & COGOLLO
FIGURE 1. Compsoneura choibo. A. Terminal branch with a female inflorescence showing two leaves on the upper surface and one
leaf on the lower surface (from B. Villanueva-T. 296). B. Staminate inflorescences (from B. Villanueva-T. 1897). C. Staminate flower. D.
Pistillate flower. E. Fruit. F. Fruit without pericarp showing the seed (lateral view). G. Fruit without pericarp showing the very short and
inconspicuous aril (basal view) (C to G from A. Montes 216). Drawn by Omar Bernal.

A new species of Compsoneura Phytotaxa 637 (3) © 2024 Magnolia Press • 253
FIGURE 2. Compsoneura choibo. A. Branch with female inflorescences and leaves on the under and upper face (B. Villanueva-T. 296)
B. Staminate flower with thinly ferruginous tomentellous perianth on the outer surface, exposing the anthers (lateral view) (B. Villanueva-
T. 1897). C. Fruit narrowly apiculate at the apex; pericarp strong, densely ferruginous pubescent, apparently indehiscent or partially
dehiscent-by-force. Seed with very short and inconspicuous aril, red-pink color when fresh, testa, bright brown with tannish red mottling
when fresh (A Montes 216). D. Inflorescence staminate. Lankester plate by Mario Sierra. Photos: A, B, D B. Villanueva-T; C, Andrés
Montes.

254 • Phytotaxa 637 (3) © 2024 Magnolia Press VILLANUEVA-TAMAYO & COGOLLO
Dioecious trees to 18 m tall, the inner bark exuding red-purple sap; wood cream colored; branchlets brown when
dry, 2.3–3.5 mm wide, glabrous, smooth and slightly striate longitudinally, when young densely brown tomentellous
(hairs minutely T-shaped). Leaves simple, alternate; petioles stout, ferruginous to light-brown when dry, 6–11.2 ×
1–1.5 mm, slightly or strongly canaliculated, minutely to densely ferruginous-tomentellous; leaf blades papery thin
to thin chartaceous, elliptic to ovate-elliptic, 10.5–15.6 × 3.5–5.5 cm, glabrous; the base acute to narrowly obtuse
or slightly attenuate; the apex acute or acuminate, acumen 4.2–6.3 mm long; the margin slightly revolute; surface
shining above, dull beneath, light-green when fresh, dull-green to green-olive when dry, resinous punctate throughout,
especially in young leaves; midrib minutely raised above, strongly raised below; secondary veins 8–9 per side, spaced
9–19.7 mm, arcuate-ascending, strongly eucamptodromous, the veins free, ascending, but not anastomosing near
margins; Staminate inflorescences 2.6–5.7 × 1.4–1.5 cm, axillary, once paniculate, with sparse T-shaped trichomes,
1–2-fasciculate; rachis 4–18 mm long, opposite first branched, alternate others, branches 2.8–6.6 mm long. Staminate
flower buds subglobose or obovoid-acuminate, staminate flowers vasiform, arranged in clusters of about 6–15 flowers
per inflorescence branch; pedicels slender, 1.3–3.5 × 0.2–0.5 mm; perianth thin coriaceous, 3.4–5.8 × 1.4–1.8 mm,
tepal lobes 3, erect, narrowly ovate to lanceolate-ovate or narrowly deltoid, 4.3–5.7 × 1–1.2 mm; perianth tube 2.3–1.7
mm long; staminate perianth vestiture thinly ferruginous tomentellous outside, simple trichomes inside; androecium
ca. 1.5 mm long, filament column 0.2–0.3 × 0.2–0.3 mm; anthers 5, fused at base for 1/3, erect, ascending from base
to apex, 1.2–1.4 × 0.2–0.3 mm, acute to apex. Pistillate inflorescences axillary, once paniculate, 2–3 branched, 8–10
× 5–7 mm, with 3–4 flowers. Pistillate flowers vasiform to cupuliform, scarcely pubescent adaxially, the hairs short
glandular, abaxially glabrescent; perianth coriaceous to carnose, 3.8–4.2 × 3.3–3.8 mm, tepal lobes 3, 2–2.4 × 0.9–1.2
mm, slightly reflexed at apex, narrowly ovate to ovate; perianth tube 1.8–2.2 mm long; perianth thick coriaceous,
vestiture glabrous inside, sparsely tomentose outside, trichomes T-shaped; ovary ovate, tannish-brown when dry,
densely ferruginous-tomentose, 0.6–0.8 × 0.8–0.9 mm, stigma bilobed, light-brown when dry, ca. 0.2 mm long, 0.2–0.4
mm wide at apex. Infructescence axillary, 12–15.2 mm long, solitary or paired; peduncle 9.4–10.2 × 4.1–4.5 mm, terete
to subterete, scarcely pubescent; fruits 1–2 per infructescence. Fruits elliptic to elliptic-ovate, 4.3–4.7 × 2.6–2.9 cm,
narrowly rostrate at the apex, rostrum 5–8 mm long; pericarp 2–3 mm thick when fresh, strong, not carinate, densely
pubescent (ferruginous tomentellous with T-shaped trichomes), apparently indehiscent or partially dehiscent-by-force.
Seeds 4.1–4.4 × 2.3–2.6 mm, aril reduced to the basal part, very short and inconspicuous, less than 5 mm long, red-pink
when fresh, tannish to red than when dry, testa bright brown with tannish red mottling; endosperm white, entire.
Distribution and habitat: This new species is presumably endemic to the middle Magdalena River and lower
Cauca River valleys. It has been collected in wet low forests, over well drained hills in the Santander and Antioquia
departments, in Colombia (Fig. 3).
Additional specimens examined: COLOMBIA. Antioquia: Zaragoza, Bajo tigüí, 200 m.a.s.l., 22 june 1967 (fl
♂), C. Yúñez 40 (UDBC!); Puerto Berrio, Vereda la Cristalina, Hacienda la Guajira, 300 m.a.s.l., 6.35000, -74.58333,
10 Sept. 2004 (fr), A. Arango et al. 402 (HUA!). Santander: Cimitarra, Bocas del Carare, Caño San Juan, Hacienda
Bocas del San Juan, Parcelas de Investigación, bosque tierra firme, 115 m.a.s.l., 6.68333, -74.11667, 24 February
2010 (fr), B. Villanueva-T. & L. Abondano 265 (TOLI!); Ibidem, 114 m.a.s.l., 6.69639, -74.11972, 17 April 2010 (fl
♀), B. Villanueva-T. & L. Abondano 296 (JBB! [accession 39802]); Ibidem, 9 Aug 2010 (fr), B. Villanueva-T. & S.
Figueroa 524 (TOLI!); Ibidem, Bosque Choibo albino, 122 m.a.s.l., 6.68806, -74.14806, 4 Dec. 2014 (fr), A. Montes
216 (TOLI!, JBB!); Sabana de Torres, Loma de Tigre, 300 m.a.s.l. (without geographic coordinates), 10 Dec. 1977 (fr),
E. Rentería et al. 28 (COL!).
Phenology:—Collected with staminate flowers in April and June, pistillate flowers in April, and with fruits in
February, August, September, and December.
Conservation status:—Compsoneura choibo has an area of occupancy (AOO) of 1200 km² and an extent of
occurrence (EOO) of approximately 4,200 km², and according to the IUCN criteria (Bachman et al. 2011), it should
be considered “Endangered” (EN) (B1ab). In addition, none of the three localities corresponds to protected areas, and
they are widely deforested for livestock farming in Santander department, and for mining and livestock farming in
Antioquia department.
Etymology:—The specific epithet refers to the primate species Ateles hybridus Geoffroy (1829: 168), known in
the region by the common name of “choibo”. This epithet is a classic case of a name in apposition according to Art.
23.1 of the International Code of Nomenclature for algae, fungi, and plants (Turland et al., 2018). The new species was
discovered in the same areas of the Middle Magdalena forests where the “choibo” monkey occurs.
Common names:—The names “soto” and “sangretoro” were recorded during fieldwork in the Antioquia
department.

A new species of Compsoneura Phytotaxa 637 (3) © 2024 Magnolia Press • 255
FIGURE 3. Geographical distribution of Compsoneura choibo in the middle Magdalena and lower Cauca river basins, Antioquia and
Santander departments, Colombia.

256 • Phytotaxa 637 (3) © 2024 Magnolia Press VILLANUEVA-TAMAYO & COGOLLO
 Natural history:—The primary knowledge of C. choibo comes from the wet forests in the interandean valleys of
the Magdalena River, specifically within the ‘Magdalena middle’ region (Hernandez 1992), a place characterized by
high rainfall and low drainage (Aldana et al. 2017). These forests represent a dynamic mosaic of várzea floodplains
and low hills terra-firme areas, which exhibit higher species richness and turnover rates (Casas et al. 2017, Umaña et
al. 2018).
In the department of Santander, near the Carare River (an affluent of Magdalena River), five permanent monitoring
plots were established in 2010 and reevaluated in 2013. Among them, three plots were located in terra-firme, and two
in várzea alluvial floodplains. Compsoneura choibo was exclusively observed within the terra-firme plots, with a total
of 10 individuals measured, exhibiting an average diameter at breast height (DBH) of ca. 13 cm (Villanueva 2020).
These forest ecosystems, although situated on flat terrain, are characterized by slow drainage and remain non-flooded
places (Villanueva 2020). This environmental condition (terra-firme) acts as a selective filter exclusive to members
of the Myristicaceae (Villanueva 2020). In this locality, this species inhabits the same forests where research has been
conducted by the ‘Primates of Colombia’ project, primarily focusing on the behavior of spider monkeys (A. hybridus),
which act as effective seed dispersers for several plant species (Suarez 2006, Gonzales et al. 2009, Link et al. 2012).
Discussion:—This new species is identifiable through a unique combination of features. Notably are its fruits
with a smooth pericarp, without carinae or sulcations, densely covered by T-shaped ferrugineous trichomes (Figs. 1A,
F, G, 2A–C) and a short (ca. 5 mm long) red-pink aril. This reduced aril was originally observed by Janovec (2000).
Regarding leaf size and the eucamptodromous venation, C. choibo resembles C. mexicana. However, C. mexicana
has leaves with a leatherier texture, pistillate flowers with erect lobes and fruits with a smooth, shiny pericarp, without
rostrum. In contrast, C. choibo has leaves papery thin to thin chartaceous, pistillate flowers with reflexed perianth
lobes, and fruit with dull pericarp with rostrum.
In addition, this new species is morphologically related to taxa that belong to the informal group known as
‘Group Hadrocarpa’ (Janovec 2000). This group is divided in two subgroups, one of which is known as ‘Complex
Compsoneura atopa’ including two species distributed in the Middle Magdalena region: Compsoneura anoriensis
and Compsoneura claroensis, and characterized by having a reduced white aril (Janovec & Neill 2003), a feature that
contrasts with the red-pink aril of C. choibo. Nonetheless, this new taxon differs from all other species of the genus in
the characters presented in the diagnosis and in the following identification key.

Key to the species of Compsoneura

Couple 14 and 15 based in Smith & Woodhouse (1937)

1a. Leaf blades secondary venation brochidodromous.............................................................................................................................2

1b. Leaf blades secondary venation eucamptodromous............................................................................................................................9
2a. Leaf blades elliptic, elliptic-ovate or obovate-elliptic, 15–35 cm long...............................................................................................3
2b. Leaf blades oblong, obovate or, if elliptic, longer than 40 cm long....................................................................................................6
3a. Fruits with a thin pericarp, < 1.5 mm thick, slightly carinate on one side, with 9–10 subsidiary longitudinal costae (irregularly
ridged); Pacific forests of Panama, Colombia, and Ecuador............................................................................................C. rigidifolia
3b. Fruits with a pericarp ≥ 2 mm thick, strongly carinate, with fewer than 9 irregular and thick costae; Andean region of Colombia,
Amazonian Ecuador, and Peru............................................................................................................................................................4
4a. Secondary venation weakly brochidodromous, the secondary veins anastomosed near the margin; fruits sessile..............................
.....................................................................................................................................................................................C. nallarettiana
4b. Secondary venation strongly brochidodromous; fruits pedicellate.....................................................................................................5
5a. Leaf blades and perianth covered by dendritic trichomes, sepals thick, extremely rigid, perianth lobes cucullate. Amazonian
Ecuador (250–400 m elev.)..............................................................................................................................................C. lapidiflora
5b. Leaf blades and perianth glabrous or glabrescent throughout, the trichomes minute, sessile or short-stalked, 2-branched, not
dendritic; sepals thin, not extremely rigid, perianth lobes erect and elongated; foothills and Andean Ecuador and Peru (600–2160
m elev.).................................................................................................................................................................................... C. diazii
6a. Leaf blades 40–50 cm long, oblong to oblong-elliptic; midrib and secondary veins covered by ferrugineous stellate trichomes on
the lower surface......................................................................................................................................................................C. atopa
6b. Leaf blades less than 35 cm long, oblong to oblong-elliptic or obovate; midrib and secondary veins glabrescent to sparsely
tomentellous on the lower surface.......................................................................................................................................................7
7a. Leaf blades obovate to orbiculate, glabrescent to ferrugineous-tomentellous when young, glabrescent to glabrous when mature on
the lower surface; trichomes minute, 2-branched, drying red brown; Colombian Pacific coast................................ C. cuatrecasasii
7b. Leaf blades oblong to oblong-elliptic, glabrescent to sparsely tomentellous on the lower surface, trichomes minute, 3–4-branched,
northern Colombian interandean valleys.............................................................................................................................................8
8a. Leaf blades drying grayish brown; fruit with longitudinal ridges giving rise to hornlike projections; Anori River valley (northern
Colombian Andes)...........................................................................................................................................................C. anoriensis
8b. Leaf blades drying brown to dark brown; fruit with longitudinal ridges, lacking hornlike projections; middle Magdalena river
valley............................................................................................................................................................................... C. claroensis

A new species of Compsoneura Phytotaxa 637 (3) © 2024 Magnolia Press • 257
9a. Fruits carinate, leaf blades sparsely tomentellous at base...............................................................................................C. capitellata
9b. Fruits non-carinate, leaf blades glabrous at base...............................................................................................................................10
10a. Leaf blades coriaceous to thick coriaceous, staminate inflorescence reduced to a tightly clustered axillary panicle, ≤ 1.3 cm long..
............................................................................................................................................................................................... C. debilis
10b. Leaf blades membranaceous to thin coriaceous, staminate inflorescence well-developed, > 1.3 cm long.......................................11
11a. Staminate inflorescence a single raceme...........................................................................................................................................12
11b. Staminate inflorescence a paniculate.................................................................................................................................................13
12a. Leaf blades membranaceous, with less than 6 secondary veins when mature................................................................. C. racemosa
12b. Leaf blades chartaceous, with more than 6 secondary veins when mature................................................................................C. ulei
13a. Staminate flower buds obovate.........................................................................................................................................................14
13b. Staminate flower buds ovate.............................................................................................................................................................16
14a. Staminate inflorescences 4–9 cm long; perianth often 4-lobed; flowers 2 or 3 per fascicle, those toward apex of inflorescence
solitary; filament column carnose or spongiose, swollen distally and leading into a carnose obconical connective; Costa Rica,
Honduras, and Panama......................................................................................................................................................... C. excelsa
14b. Staminate inflorescences 2–4 cm long; perianth 3-lobed; flowers 3–7 per fascicle; filament column not carnose or spongiose, not
swollen distally; Colombia................................................................................................................................................................15
15a. Leaf blades papyraceous, usually translucent, slightly obovate, the apex 10–15 mm long, abruptly acuminate; Colombian
interandean valleys................................................................................................................................................................C. mutisii
15b. Leaf blades thick coriaceous, opaque, ovate, ovate-elliptic or elliptic, the apex ca. 5 mm long, cuspidate; Colombian Pacific
coast...................................................................................................................................................................................... C. trianae
16a. Fruit densely pubescent, ferrugineous or ferrugineous-tomentellous, the pericarp strong and ligneous..............................C. choibo
16b. Fruit glabrous, the pericarp thin, non-ligneous.................................................................................................................................17
17a. Shrubs, treelets or small trees; leaf blades thick coriaceous, strongly translucent; Colombian Amazon basin ........ C. schultesiana
17b. Trees; leaf blades chartaceous or papyraceous, slightly translucent.................................................................................................19
18a. Perianth vasiform, lobes erect; leaf blades elliptic, elliptic-obovate; stem tan, striate, bark not exfoliating; Central America...........
.......................................................................................................................................................................................... C. mexicana
18b. Perianth cupuliform, urceolate, lobes recurved at apex; leaf blades elliptic to elliptic-ovate; stem brown or darker black, bark
exfoliating; Amazon basin of Brazil, Colombia, Ecuador, Peru, and Venezuela................................................................. C. sprucei

Acknowledgements

The authors are grateful to the Primates of Colombia Foundation for their support in making this discovery possible,
with special thanks to A. Link, A. Montes, and their dedicated field support team. The first author acknowledged
the University of Tolima for a research grant, and appreciates O. Bernal and M. Sierra for their contributions to the
illustration and Lankester plate. The authors also thank A. Orejuela (JJB), D. Santamaría-Aguilar and G. Aymard
(PORT) for their valuable comments and insights to the manuscrit. To K. N. Gandhi (GH) for his nomenclatural
expertise. Our gratitude to curators and directors to the following herbaria: BOG, COAH, COL, CUVC, FMB, HPUJ,
HUA, JBB, MEDEL, TOLI, and UDBC. who allowed access to the collections.

References

Aldana, A.M., Villanueva, B., Cano, A., Correa, D., Umaña, M.N., Casas, L.F., Cárdenas, C., Henao-Diaz, L.F. & Stevenson, P.R. (2017)
Drivers of biomass stocks in Northwestern South American forests: Contributing new information on the Neotropics. Forest Ecology
and Management 389: 86–95.
https://doi.org/10.1016/j.foreco.2016.12.023
Aymard, G.A., Castro-Lima, F. & Arellano-P, H. (2020) Identificación de Myristicaceae de la cuenca del Río Negro en ausencia de flores
y frutos. Pittieria 44: 28–55.
Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: geospatial
conservation assessment tool. ZooKeys 150: 117–126.
https://doi.org/10.3897/zookeys.150.2109
Brown, R. (1810) Prodromus florae Novae Hollandiae et Insulae Van-Diemen, exhibens characteres plantarum. R. Taylor, Londres, 464
pp. [https://www.biodiversitylibrary.org/page/2954555]
Candolle, A.L.P.P. de. (1856) Prodromus Systematis Naturalis Regni Vegetabilis, v.14. Sumptibus Sociorum Treuttel et Würtz, Paris, 718
pp. [https://www.biodiversitylibrary.org/page/35138406]
Casas, L.F., Aldana, A.M., Henao-Diaz, F., Villanueva, B. & Stevenson, P.R. (2017) Specific gravity of woody tissue from lowland
Neotropical plants: differences among forest types. Ecology 98 (5): 1474.

258 • Phytotaxa 637 (3) © 2024 Magnolia Press VILLANUEVA-TAMAYO & COGOLLO
 https://doi.org/10.1002/ecy.1786
Gentry, A.H. (1993) A field guide to the families and genera of woody plants of northwest South America (Colombia, Ecuador, Perú), with
supplementary notes on herbaceous taxa, Conservation International, Washington D.C., 895 pp.
Geoffroy, I. (1829) Description de Deux Especes Nouvelles de Singes a Queue Prenante (Ateles hybridus). Mémoires du Muséum national
d’Histoire naturelle 17: 168–173. [https://www.biodiversitylibrary.org/page/35009806]
González-Zamora, A., Arroyo-Rodríguez, V., Chaves, O.M., Sánchez-López, S., Stoner, K.E. & Riba-Hernández, P. (2009) Diet of spider
monkeys (Ateles geoffroyi) in Mesoamerica: current knowledge and future directions. American Journal of Primatology 71: 8–20.
https://doi.org/10.1002/ajp.20625
Gronovius, J.F. (1755) Flora Orientalis sive Recensio Plantarum, Wilhelmi de Groot, Leiden, 175 pp.
Hallé, F., Oldeman, R.A.A. & Tomlinson, P.B. (1978) Tropical trees and forests: an architectural analysis, Springer-Verlag, New York,
385 pp.
https://doi.org/10.1007/978-3-642-81190-6
Hernández, J. (1992) Caracterización geográfica de Colombia. In Halffter, G. (Ed.) La diversidad biológica de Iberoamérica I. Acta
Zoológica Mexicana, Xalapa, pp. 105–1552.
Jaramillo, T.S. & Balslev, H. (2001) Two new Myristicaceae from Ecuador. Nordic Journal of Botany 21: 561–566.
https://doi.org/10.1111/j.1756-1051.2001.tb00809.x
Janovec, P.J. (2000) A systematic study of Compsoneura (Candolle) Warburg, a Neotropical member of the nutmeg family (Myristicaceae)
PhD. thesis, Texas A&M University, College Station, Texas, 359 pp.
Janovec, J.P. & Harrison, J.S. (2002) A Morphological Analysis of the Compsoneura sprucei Complex (Myristicaceae), with a New
Combination for the Central American Species Compsoneura mexicana. Systematic Botany 27 (4): 662–673. Available from: http://
www.jstor.org/stable/3093914 (accessed: 21 February 2024).
Janovec, J.P. & Neill, A.K. (2003) Studies of the Myristicaceae: An Overview of the Compsoneura atopa Complex, with Descriptions of
New Species from Colombia. Brittonia 54 (4): 251–261.
https://doi.org/10.1663/0007-196X(2003)54[251:SOTMAO]2.0.CO;2
Janovec, J.P. (2002) Compsoneura diazii (Myristicaceae), a New Species from the Rio Cenepa Area of Northwestern Peru. Novon 12 (3):
366–368.
https://doi.org/10.2307/3393080
Kühn, U. & Kubitzki, K. (1993) Myristicaceae. In: Kubitzki, K., Rohwer, J.-G. & Bittrich, V. (Eds.) The Families and Genera of Vascular
Plants. Flowering Plants. Dicotyledons. Magnoliid, Hamamelid, and Caryophyllid families. Springer Verlag, Berlin, pp. 457–467.
https://doi.org/10.1007/978-3-662-02899-5_53
Link, A., Galvis, N., Marquez, M., Guerrero, J., Solano, C. & Stevenson, P.R. (2012) Diet of the critically endangered brown spider
monkey (Ateles hybridus) in an inter—Andean lowland rainforest in Colombia. American Journal of Primatology 12: 1097–1105.
https://doi.org/10.1002/ajp.22066
Markgraf, F. (1926) Plantae Tessmannianae Perúvianae III. Notizblatt Des Königl. Botanischen Gartens Und Museums Zu Berlin 9 (89):
964.
https://doi.org/10.2307/3994448
Rios, P., Zarate, R. & Grandes-Rios, J. (2023) Compsoneura nallarettiana (Myristicaceae), a new species from north-western Perú.
Phytotaxa 592: 294–300.
https://doi.org/10.11646/phytotaxa.592.3.7
Rodrigues, W.A. (1989) Two new Neotropical species of Compsoneura (Myristicaceae). Brittonia 41 (2): 160–163.
https://doi.org/10.2307/2807526
Rodrigues, W.A., Aymard, G. & Berry, P. (2001) Myristicaceae. In: Berry, P., Yatskievych, K. & Holst, B.K. (Eds.) Flora of the Venezuelan
Guayana. Vol. 6. Missouri Botanical Garden Press, St. Louis, pp. 734–751.
Santamaría-Aguilar, D. & Lagomarsino, L.P. (2022) New Species of Virola (Myristicaceae) from South America. Phytotaxa 197: 81–
148.
https://doi.org/10.3897/phytokeys.197.81367
Smith, A.C. & with the collaboration of Woodhouse, R.P. (1937) The American species of Myristicaceae. Brittonia 2: 393–510.
https://doi.org/10.2307/2804799
Smith, A.C. (1938) Compsoneura. Brittonia 2 (5): 405–416.
https://doi.org/10.2307/2804801
Smith, A.C. (1950) Studies of South American plants. XII. Contributions from the United States National Herbarium 29 (8): 318–320.
Smith, A.C. (1956) Studies of South American plants. XV. American Journal of Botany 43 (8): 573–574.
https://doi.org/10.1002/j.1537-2197.1956.tb10536.x
Spruce, R. (1857) Addenda et Corrigenda, Prodromus Systematis Naturalis Regni Vegetabilis 14: 696.

A new species of Compsoneura Phytotaxa 637 (3) © 2024 Magnolia Press • 259
 https://doi.org/10.1515/9783112430682-007
Suarez, S.A. (2006) Diet and Travel Costs for Spider Monkeys in a Nonseasonal, Hyperdiverse Environment. International Journal of
Primatology 27: 411–436.
https://doi.org/10.1007/s10764-006-9023-6
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold,
K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (eds.) (2018) International Code of Nomenclature for
algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017.
Regnum Vegetabile 159. Glashütten, 322 pp.
https://doi.org/10.12705/Code.2018
Umaña, M.N., Correa, D., Cano, A., Casas, L.F., Cárdenas, S., Villanueva, B. & Stevenson, P. (2018) Diversity of Dispersal Systems in
Igapó Forests: An Analysis of Local Tree Diversity, Species Turnover, and Dispersal Systems. In: Myster, R. (Eds.) Igapó (Black-
water flooded forests) of the Amazon Basin. Springer, Switzerland, pp. 23–35.
https://doi.org/10.1007/978-3-319-90122-0_2
Villanueva, B. (2020) Diversidad y dinámica del carbono en bosques húmedos tropicales de tierras bajas, ubicados en la cuenca inferior
del río Carare, Cimitarra, Santander. M.Sc. thesis, Universidad del Tolima. Ibagué, 169 pp.
Warburg, O. (1897) Monographie der Myrisiticaceen. Nova acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae
Curiosorum 68: 1–146.
Warburg, O. (1905) XXVI. Myristicaceae Costaricenses. Repertorium Specierum Novarum Regni Vegetabilis 1: 71–72.
https://doi.org/10.1002/fedr.19050010503

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