A new species in the tree genus Polyceratocarpus (Annonaceae) from the Udzungwa Mountains of Tanzania
Author
Marshall, Andrew R.
CIRCLE, Environment Department, University of York, York, UK & Flamingo Land Ltd., Kirby Misperton, North Yorkshire, UK
andy.marshall@york.ac.uk
Author
Couvreur, Thomas L. P.
https://orcid.org/0000-0002-8509-6587
Institut de Recherche pour le Developpement, UMR-DIADE, BP 64501, F- 34394 Montpellier cedex 5, France & Universite de Yaounde I, Ecole Normale Superieure, Departement des Sciences Biologiques, Laboratoire de Botanique systematique et d'Ecologie, B. P. 047, Yaounde, Cameroon & Netherlands Centre for Biodiversity Naturalis (section NHN), Biosystematics Group, Wageningen University, the Netherlands
Author
Summers, Abigail L.
CIRCLE, Environment Department, University of York, York, UK & Flamingo Land Ltd., Kirby Misperton, North Yorkshire, UK & Biology Department, University of York, York, UK
Author
Deere, Nicolas J.
CIRCLE, Environment Department, University of York, York, UK & Durrell Institute of Conservation and Ecology, School of Anthropology and Conservation, University of Kent, Canterbury, UK
Author
Luke, W. R. Quentin
East African Herbarium, National Museums of Kenya, Nairobi, Kenya
Author
Ndangalasi, Henry J.
Department of Botany, University of Dar es Salaam, Dar es Salaam, Tanzania
Author
Sparrow, Sue
Biology Department, University of York, York, UK
Author
Johnson, David M.
Department of Botany & Microbiology, Ohio Wesleyan University, Delaware, Ohio 43015 USA
text
PhytoKeys
2016
2016-06-01
63
63
76
http://dx.doi.org/10.3897/phytokeys.63.6262
journal article
http://dx.doi.org/10.3897/phytokeys.63.6262
1314-2003-63-63
C12BFF8EBE10FFD42532FFA2FFC05F36
899007
Polyceratocarpus askhambryan-iringae A.R. Marshall & D.M. Johnson
sp. nov.
Figs 1
, 2
, 3
Diagnosis.
This species may be distinguished from other species of
Polyceratocarpus
by the combination of glabrous non-glaucous leaves with finely reticulate to weakly scalariform tertiary venation, pedicels 15-22 mm long, broadly ovoid buds, chartaceous petals 10-17 mm long, 5 to 18 carpels/monocarps, and relatively large torulose monocarps.
Type
.
Marshall 2117
(
holotype
K; isotypes DSM, MO, NHT),
Tanzania
,
Iringa Region
:
Ndundulu Forest
,
Kilombero Nature Reserve
,
Udzungwa Mountains
,
07°48'S
,
36°31'E
(WGS84),
1490 m
,
30 May 2011
.
Description.
Monopodial
tree
to 20 m tall, 4.0-25.4 cm diam.; bark smooth, sparsely lenticellate, often with weak horizontal striations and pits on large trees, grey-brown; branches spirally arranged on trunk, branching from half to two fifths of the height of the main stem, perpendicular but sinuous and drooping slightly; twigs longitudinally rugulose, inconsistently marked with small but prominent lenticels, glabrous, brown.
Leaves
: petiole 4-9 mm long, 1.6-3.3 mm thick, roughened, black, glabrous; lamina narrowly to broadly elliptic-oblong to oblanceolate, or rarely obovate, (5.0-)9.0-25.7 by (3.7-)4.9-8.6(-11.6) cm, chartaceous to coriaceous, greenish gray in sicco, glabrous on both surfaces, base rounded and minutely subcordate, apex acuminate with the acumen 12-20 mm long or occasionally obtuse, midvein plane to slightly impressed above, raised below, secondary veins 9 to 17 per side, diverging at 45-60° from midrib, eucamptodromous to weakly brochidodromous, slightly raised to slightly impressed above, raised below, tertiary veins finely reticulate to somewhat scalariform, indistinct to slightly raised above, raised and conspicuous below.
Inflorescences
1- or 2-flowered, axillary or occasionally ramiflorous, forming tubercles on leafless growth; pedicels 15-29 mm long 1-3 mm diam., finely appressed-puberulent, bearing a bract 0.8-1 mm long 1/4-2/5 of the distance above pedicel base. Flowers bisexual or staminate, buds broadly ovoid; sepals 3, valvate, crescent-shaped, 2-3.5 mm long, partially connate at the base so that as corolla expands the calyx becomes discoid to triangular with diameter of 7-8 mm, appressed-puberulent abaxially; petals in two whorls of 3, pale yellow in vivo; outer petals occasionally tinged pink on abaxial surface in vivo, valvate, spreading horizontally and recurving at anthesis, narrowly elliptic to elliptic or oblong-elliptic, 10-16 mm long by (5-)8-11 mm wide, coriaceous, apex obtuse, sparsely pubescent adaxially, ferruginous appressed-puberulent but becoming glabrate and verrucose abaxially; inner petals sometimes with a pale brownish-yellow median stripe abaxially, valvate, erect at anthesis with the apices recurved, narrowly elliptic-lanceolate, elliptic, or elliptic-oblanceolate, 11-17 mm long
x
5-7 mm wide, coriaceous, external surface marked by a broad flattened ridge that narrows from base to apex, glabrous adaxially, appressed-puberulent with
trichomes
densest along ridge and at apex abaxially, verrucose on both surfaces, apex acute; stamens ca. 200, 2.0-2.8 mm long, clavate, apex of connective obliquely truncate, pale brown with orange apex, glabrous?; carpels 5 to 18, oblong, 2.9-4.0(-6.0) mm long by 0.9-1.1(-2.3) mm wide, densely pale brown/ferruginous-puberulous; stigma bilobed, capitate, 1 mm in diam., glabrous, ovules ca. 10, uniseriate; torus subglobose to broadly pyriform to oblate, 4.4-4.9 mm long by 2.6-5.4 mm diam., 3-8 mm diam. at base. Pedicel of fruit 20-44 mm long by 3-7 mm diam., weakly longitudinally rugulose, glabrate; torus of fruit ellipsoid to broadly pyriform, 7-15 mm diam.
x
8-12 mm long, grey-brown.
Monocarps
up to 18 per fruit, green (rarely with orange or vinaceous tinge) in vivo, dark brown when dried, weakly (to strongly) recurved-falciform, (1.9) 6.0-8.6 cm by 0.7-2.2 cm, torulose, minutely verrucose, glabrate or with a few scattered hairs, base sub-sessile or short-stipitate, stipe 1-11 mm long, 2-6 mm thick, apex rounded or sometimes short-beaked.
Seeds
1-15 per monocarp, 15 mm long by 13 mm wide by 10 mm thick, arranged in a single [or two irregular?] rows, flattened-ellipsoid, pitted, with spiniform ruminations (fig. 3) and raphe/antiraphe sunken in a circumferential groove.
Distribution.
Polyceratocarpus askhambryan-iringae
is endemic to the Udzungwa Mountains of Tanzania. It is known from Mwanihana Forest in the Udzungwa Mountains National Park, Ndundulu Forest in the Kilombero Nature Reserve, and the Uzungwa Scarp Forest Reserve (Fig.
1
).
Figure 1.
Known distribution of
Polyceratocarpus askhambryan-iringae
in the Eastern Arc Mountains (EAM). EAM boundary and forest cover (green) derived from
Platts et al. (2011)
. Black boundaries within EAM boundary show protected areas. Pale green areas in the lower maps show degraded forest with canopy <10%.
Figure 2.
Polyceratocarpus askhambryan-iringae
drawings of
A
tree architecture
B
fresh fruits
C
fresh flower below
D
fresh ramiflorous flower buds
E-F
dry and fresh bisexual flower (one petal removed)
G
fresh bisexual flower above
H
dried stamens
I
-
J
fresh and dry carpels lacking stigmas
K
dried carpel with stigma, plus photographs of
L
fresh leaves
M
fruit and
N
flower. Drawings by Sue Sparrow,
A
by Andrew Marshall,
E
and
K
by Andrew Brown, from the following specimens: Marshall 2070 (
B
); Marshall 2117 (
C-E
and
G-L
) and Luke 11279 (
F
). Scale bars: 20 mm unless stated.
Figure 3.
SEM photograph of
Polyceratocarpus askhambryan-iringae
dried seed cross-section, showing spiniform ruminations.
Habitat and ecology.
Inhabits montane forest on brown sandy loam soils (pH range 4-5 measured in Ndundulu Forest). Mean annual rainfall of collection localities approximately 1500-2000 mm/yr (Marshall, Ndangalasi, unpubl. data). Thirty-eight mature individuals were found mostly on slopes or ridge-tops at elevations 1090-1540m. Mature flowers were collected in May, November, and December, fruits in February and May-October.
Associated taxa recorded with
Polyceratocarpus askhambryan-iringae
include the following:
(1) Ndundulu Forest
:
Allanblackia ulugurensis
Engl.,
Alsodeiopsis schumannii
Engl.;
Anisotes pubinervius
(T.Anderson) Heine,
Anthocleista grandiflora
Gilg,
Beilschmiedia kweo
(Mildbr.) Robyns & Wilczek,
Bertiera pauloi
Verdc.,
Cassipourea gummiflua
Tul.;
Cassipourea malosana
Alston;
Chlorophytum brachystachyum
Baker,
Cleistanthus polystachyus
Hook.f. ex Planch.,
Clerodendrum cephalanthum
Oliv.,
Coffea
sp.,
Cola greenwayi
Brenan,
Cola stelechantha
Brenan;
Craterispermum longipedunculatum
Verdc.;
Diospyros abyssinica
(Hiern) F.White,
Drypetes gerrardii
Hutch.;
Englerina
sp. nov.;
Garcinia buchananii
Baker;
Garcinia volkensii
Engl.;
Grewia mildbraedii
Burret;
Justicia rodgersii
Vollesen;
Lasiodiscus usambarensis
Engl.;
Maytenus undata
(Thunb.) Blakelock;
Monodora globiflora
Couvreur;
Monanthotaxis schweinfurthii
Engl. & Diels;
Myrianthus holstii
Engl.,
Ocotea usambarensis
Engl.;
Ochna holstii
Engl.;
Parinari excelsa
Sabine,
Peddiea fischeri
Engl.,
Pavetta nitidissima
Bridson,
Plectranthus leptophyllus
(Baker) A.J.Paton,
Rinorea angustifolia Baill. subsp. ardisiiflora
(Oliv.) Grey-Wilson;
Rinorea
sp.;
Rytigynia lichenoxenos (K.Schum.) Robyns subsp. glabrituba
Verdc.;
Sclerochiton obtusisepalus
C.B.Clarke;
Solanecio epidendricus
(
Mattf.) C.Jeffrey;
Strombosia scheffleri
Engl.;
Strychnos mellodora
S.Moore;
Strychnos mitis
S.Moore;
Strychnos
sp. nov;
Syzygium guineense
DC.;
Tabernaemontana stapfiana
Britten;
Tarenna pavettoides
(Harv.) Sim;
Thalictrum rhynchocarpum
Quart.-Dill. & A.Rich.;
Uvariopsis lovettiana
Couvreur & Q.Luke;
Vepris stolzii
I.Verd.;
Vernonia calvoana Engl. subsp. leucocalyx
(O.Hoffm.) C.Jeffrey;
Vernonia luhomeroensis
Q.Luke & Beentje;
Vernonia sp. nr. pteropoda
Oliv. & Hiern;
Warneckea
sp. nov.;
Xymalos monospora
Baill.;
Zanthoxylum paracanthum
(Mildbr.) Kokwaro;
Zehneria sp. nr. oligosperma
C.Jeffrey.
(2) Uzungwa Scarp Forest Reserve
:
Cassipourea gummiflua
Tul;
Cleistanthus polystachyus
Hook.f. ex Planch.;
Craterispermum longipedunculatum
Verdc.;
Diospyros uzungwaensis
Frim.-
Moll
. & H.J.Ndangalasi;
Drypetes gerrardii
Hutch.;
Lasiodiscus usambarensis
Engl.;
Psychotria megalopus
Verdc.;
Tabernaemontana stapfiana
Britten;
Tarenna uzungwaensis
Bridson.
(3) Mwanihana
:
Acalypha psilostachya Hochst. ex A.Rich. var. psilostachya
;
Anisotes pubinervius
(T.Anderson) Heine;
Caloncoba welwitschii
Gilg;
Chrysophyllum gorungosanum
Engl.;
Coffea mufindiensis Hutch. ex Bridson subsp. mufindiensis
;
Dorstenia sp. aff tenuiradiata
Mildbr.;
Isoglossa lactea Lindau ex Engl. subsp. lactea
;
Isolona linearis
Couvreur;
Newtonia buchananii
(Baker) G.C.C.Gilbert & Boutique;
Ochna holstii
Engl.;
Parinari excelsa
Sabine;
Phyllopentas ulugurica
(Verdc.)
Karehed
& B. Bremer;
Kedrostis
sp.;
Polystachya sp aff. canaliculata
Summerh.;
Raphidiocystis chrysocoma
(Schumach.) C.Jeffrey;
Selaginella kraussiana
(Kunze) A.Braun;
Stellaria mannii
Hook.f.;
Tricalysia aciculiflora
Robbr.;
Uvariopsis lovettiana
Couvreur & Q.Luke;
Vepris nobilis
(Delile) Mziray;
Zanthoxylum paracanthum
(Mildbr.) Kokwaro.
Additional specimens examined.
TANZANIA
.
Iringa Region
,
Kilolo District
:
east Udzungwa National Park
, forest south of
Mwanihana
hill c.
2 km
S of last camping site of
Mwanihana
trail,
1400 m
,
07°48'S
,
36°49'E
, Couvreur 101 (DSM, OWU, WAG);
Mwanihana Forest
above
Sanje village
,
1220 m
, no grid reference, Lovett 222 (K);
Udzungwa Mountains National Park
,
1200 m
,
07°48'S
,
36°49'E
, Luke 7738 (EA, K);
Udzungwa Mountains National Park
,
1440 m
,
07°42'S
,
36°52'E
, Luke 11279 (EA, NHT, MO, K);
Ndundulu FR
, Camp 589-Camp 590,
07°47'S
,
36°29'E
,
1440 m
, Luke et al. 10366 (MO);
Kilombero Nature Reserve
,
Ndundulu Forest
,
1540 m
,
07°48'S
,
36°31'E
(WGS84), Marshall 2036, 2070, (NHT, MO, K);
Uzungwa Scarp Forest Reserve
, Uluti,
1534 m
,
08°14'S
,
36°01'E
, Ndangalasi HJN 392 (DSM, OWU);
Uzungwa Scarp Forest Reserve
, Ilutila,
1709 m
,
08°13'S
,
36°01'E
, Ndangalasi 393 (DSM, OWU); Udzungwa,
Kilombero FR
,
W of Ruaha River
,
1700 m
,
Rogers
&
Hall
2300 (K);
Mwanihana Forest
above
Sanje village
,
1400 m
,
07°50'S
,
36°49'E
, Thomas 3656 (MO);
Mwanihana Forest
above
Sanje village
,
1400 m
,
07°50'S
,
36°49'E
, Thomas 3698 (MO, WAG)
.
Morogoro Region
,
Kilombero District
:
Sonjo-Mwanihana
trail,
1090 m
,
07°48'S
,
36°51'E
, Luke 5051 (EA, K)
.
An additional specimen from
Iringa Region
(
Nyambanitu Forest
, Ede 65, K), may also represent this species but bears only an old fruit pedicel lacking monocarps. Further potential
Polyceratocarpus
collections from
Iringa Region
(
Lulanda Forest Reserve
: Gereau 2651, 2664, 2665, MO; Lovett 2256, MO, WAG;
Luke
&
Luke
12779, EA & K), were identified as neither
Polyceratocarpus askhambryan-iringae
nor
Polyceratocarpus scheffleri
, while another
from
Morogoro Region
was not considered to be from this genus at all (
Kimboza Forest Reserve
: Parry 1816, TFD; cited
Verdcourt 1971
)
.
Additional field notes.
Slash
dry, slightly stringy, pale yellow (to pale peach), occasionally streaked yellow-brown, dark brown at outer edge formed by the colour of the inner bark, potpourri aroma.
Leaf
lamina dark green above, mid-green with greyish tinge below, turning greenish-grey when dried, new flush pinkish; petiole initially pale green in vivo, becoming roughened grey-brown with age; midrib yellow-green above and below in vivo.
Flower
buds broadly ovoid, green or pale brown-yellow with occasional pink tinge at apex in vivo; sepals yellowish-green in vivo.
Etymology.
This new species of
Polyceratocarpus
was named by Askham Bryan College and Iringa International School as part of a rainforest education program.
Conservation status.
Our IUCN Red List assessment for
Polyceratocarpus askhambryan-iringae
was based on "area of occupancy" (AOO), "extent of occurrence" (EOO;
IUCN 2012
) and the level of threat. The 38 observed mature
Polyceratocarpus askhambryan-iringae
stems were found at four collection localities, with only 112 km between the most distant individuals. The four collection localities represented three different levels of governmental protection, including Forest Reserve (Uzungwa Scarp), Nature Reserve (Kilombero, KNR; and also Uzungwa Scarp proposed status) and National Park (Udzungwa Mountains; UMNP), with only UMNP having the maximum level of protection under Tanzanian law (IUCN category II;
Dudley 2008
). At the time of survey, the greatest threat to
Polyceratocarpus askhambryan-iringae
was widespread timber-felling observed in Uzungwa Scarp Forest Reserve. Conversely, threats to
Polyceratocarpus askhambryan-iringae
in KNR and UMNP were much lower, both comprising only very occasional removal of pole-sized trees by villagers, and potential damage from an increasing elephant population (
Marshall et al. 2012
). KNR was further threatened by a lack of formal ranger patrols and rapid population expansion in the nearest villages. Given that the sources of threat were closely associated with the three protected areas, for the purpose of Red List assessment we considered three rather than four threat-defined
"locations"
(
IUCN 2012
). In calculating AOO, we used 10 km2 grid cells so that the projected area was
not
extrapolated far beyond the expected habitat tolerance of the species. We estimated an AOO of 300 km2 and an EOO of 1,410 km2, with EOO, AOO, habitat quality and the number of mature individuals, all presumed declining as a result of timber-felling in USFR. Given this continuing decline, plus an EOO of less than 5,000 km2, an AOO of less than 500 km2, and a population at no more than five locations, the species qualified firmly as endangered on the IUCN Red List (
IUCN 2012
), EN B1ab(i,ii,iii,v)+2ab(i,ii,iii,v). Within the Udzungwa Mountains, closed-canopy forest was not extensive at elevations suitable for
Polyceratocarpus askhambryan-iringae
, and hence we expect that future expeditions will not expand the AOO or EOO of
Polyceratocarpus askhambryan-iringae
above the IUCN endangered threshold. However, the Rubeho and Mahenge mountain blocs adjacent to Udzungwa were more poorly known, and may contain suitable habitat in which this species might also be found.