Figures (3)  Tables (7)
    • Figure 1. 

      Location map of 511 wild cacao collections in Peru. Using R software.

    • Figure 2. 

      Biplot resulting from the principal components analysis based on quantitative traits with the most contribution to the phenotypic diversity of cacao in Peruvian river basins.

    • Figure 3. 

      Dendrogram resulting from the cluster analysis based on 27 agronomic traits of 511 wild cacao accessions from Peruvian river basins.

    • Department River basin, collection and planted year Range of geographical coordinates Altitude m.a.s.l. Accessions
      Latitude south Longitude west
      Loreto
      (n = 282)
      Aypena (n = 22)
      2008
      5.15322°−5.19753° 76.01758°−76.08981° 120−133 AYP-1, AYP-2, AYP-3, AYP-4, AYP-5, AYP-6, AYP-7, AYP-8, AYP-9, AYP-10, AYP-11, AYP-12, AYP-13, AYP-14, AYP-15, AYP-16, AYP-17, AYP-18, AYP-19, AYP-20, AYP-21, AYP-22
      Marañón (n = 22)
      2008
      5.00139°−5.00222° 75.85292°−75.85792° 124−139 MACH-23, MACH-24, MACH-25, MACH-26, MACH-27, MACH-28, MACH-29, MACH-30, MACH-31, MACH-32, MACH-33, MACH-34, MACH-35, MACH-36, MACH-37, MACH-38, MACH-39, MACH-40, MACH-41, MACH-42, MACH-43, MACH-44
      Ungurahui (n = 38)
      2008
      4.76397°−4.77833° 76.44006°−76.44311° 127−144 UNG-45, UNG-46, UNG-47, UNG-48, UNG-49, UNG-50, UNG-51, UNG-52, UNG-53, UNG-54, UNG-55, UNG-56, UNG-57, UNG-58, UNG-59, UNG-60, UNG-61, UNG-62, UNG-63, UNG-64, UNG-65, UNG-66, UNG-67, UNG-68, UNG-69, UNG-70, UNG-71, UNG-72, UNG-73, UNG-74, UNG-75, UNG-76, UNG-77, UNG-78, UNG-79, UNG-80, UNG-81, UNG-82
      Pastaza (n = 24)
      2008
      4.83819°−4.86897° 76.42342°−76.43353° 129−140 PAS-83, PAS-84, PAS-85, PAS-86, PAS-87, PAS-88, PAS-89, PAS-90, PAS-91, PAS-92, PAS-93, PAS-94, PAS-95, PAS-96, PAS-97, PAS-98, PAS-99, PAS-100, PAS-101, PAS-102, PAS-103, PAS-104, PAS-105, PAS-106
      Ungumayo (n = 26)
      2008
      4.73297°−4.90708° 76.14847°−76.24644° 127−141 UGU-107, UGU-108, UGU-109, UGU-110, UGU-111, UGU-112, UGU-113, UGU-114, UGU-115, UGU-116, UGU-117, UGU-118, UGU-119, UGU-120, UGU-121, UGU-122, UGU-123, UGU-124, UGU-125, UGU-126, UGU-127, UGU-128, UGU-129, UGU-130, UGU-131, UGU-132
      Nucuray (n = 25)
      2008
      4.86900°−4.91703° 75.74594°−75.77547° 115−131 NUC-133, NUC-134, NUC-135, NUC-136, NUC-137, NUC-138, NUC-139, NUC-140, NUC-141, NUC-142, NUC-143, NUC-144, NUC-145, NUC-146, NUC-147, NUC-148, NUC-149, NUC-150, NUC-151, NUC-152, NUC-153, NUC-154, NUC-155, NUC-156, NUC-157
      Urituyacu (n = 33)
      2008
      4.80644°−4.83300° 75.39525°−75.47986° 97−130 URI-158, URI-159, URI-160, URI-161, URI-162, URI-163, URI-164, URI-165, URI-166, URI-167, URI-168, URI-169, URI-170, URI-171, URI-172, URI-173, URI-174, URI-175, URI-176, URI-177, URI-178, URI-179, URI-180, URI-181, URI-182, URI-183, URI-184, URI-185, URI-186, URI-187, URI-188, URI-189, URI-190
      Morona (N = 50)
      2009
      3.56269°−4.71822° 77.08089°−77.49225° 136−196 MOR-192, MOR-194, MOR-196, MOR-198, MOR-200, MOR-202, MOR-204, MOR-206, MOR-208, MOR-210, MOR-212, MOR-214, MOR-216, MOR-218, MOR-220, MOR-222, MOR-224, MOR-226, MOR-228, MOR-230, MOR-232, MOR-234, MOR-236, MOR-238, MOR-240, MOR-242, MOR 244, MOR-246, MOR-248, MOR-250, MOR-252, MOR-254, MOR-256, MOR-258, MOR-260, MOR-262, MOR 264, MOR-266, MOR-268, MOR-270, MOR-272, MOR-274, MOR-276, MOR-278, MOR-280, MOR-282, MOR-284, MOR-286, MOR-288, MOR-290
      Chambira (n = 20)
      2013
      4.29673°−4.52512° 74.11835°−75.09407° 113−118 CH-267, CH-269, CH-271, CH-273, CH-275, CH-277, CH-279, CH-281, CH-283, CH-285, CH-287, CH-289, CH-291, CH-292, CH-293, CH-294, CH-295, CH-296, CH-297, CH-298
      Tigre (n = 22)
      2013
      3.84153°−4.44577° 74.11407°−74.41777° 99−125 TIG-299, TIG-300, TIG-301, TIG-302, TIG-303, TIG-304, TIG-305, TIG-306, TIG-307, TIG-308, TIG-309, TIG-310, TIG-311, TIG-312, TIG-313, TIG-314, TIG-315, TIG-316, TIG-317, TIG-318, TIG-319, TIG-320
      Loreto
      (n = 80)
      Napo (n = 22)
      2013
      2.81950°−3.48998° 72.64783°−73.53973° 92−104 NAP-321, NAP-322, NAP-323, NAP-324, NAP-325, NAP-326, NAP-327, NAP-328, NAP-329, NAP-330, NAP-331, NAP-332, NAP-333, NAP-334, NAP-335, NAP-336, NAP-337, NAP-338, NAP-339, NAP-340, NAP-341, NAP-342
      Putumayo (n = 18)
      2015
      0.07867°−2.33301° 73.03614°−75.24604° 40−222 PUT(A)-460, PUT(B)-460, PUT-462, PUT-464, PUT-466, PUT-466(A), PUT-466(B), PUT-468, PUT-468(A), PUT-470, PUT-472, PUT-474, PUT-476, PUT-478, PUT-480, PUT-482, PUT-484, PUT-486
      Yavari (n = 23)
      2015
      3.88507°−4.17261 70.35288°−70.57453° 91−101 YAV-435, YAV-437, YAV-439, YAV- 441, YAV-443, YAV-445, YAV-447, YAV-449, YAV-451, YAV-453, YAV-455, YAV-457, YAV-459, YAV-461, YAV-463, YAV-465, YAV-467, YAV-469, YAV-471, YAV-473, YAV-475, YAV-477, YAV-479
      Amazonas (n = 17)
      2015
      3.85979°−4.00796° 70.53532°−71.15834° 77−101 AMZ-436, AMZ-438, AMZ-440, AMZ-442, AMZ-446, AMZ-448, AMZ-452, AMZ-454, AMZ-456, AMZ-481, AMZ-483, AMZ-485, AMZ-487, AMZ-489, AMZ-491, AMZ-493, AMZ-495
      Amazonas
      (n = 38)
      Santiago(n = 38)
      2009
      3.10756°−4.44411° 77.56778°−77.87917° 148−252 SAN-191, SAN-193, SAN-195, SAN-197, SAN-199, SAN-201, SAN-203, SAN-205, SAN-207, SAN-209, SAN-211, SAN-213, SAN-215, SAN-217, SAN-219, SAN-221, SAN-223, SAN-225, SAN-227, SAN-229, SAN-231, SAN-233, SAN-235, SAN-237, SAN-239, SAN-241, SAN-243, SAN-245, SAN-247, SAN-249, SAN-251, SAN-253, SAN-255, SAN-257, SAN-259, SAN-261, SAN-263, SAN-265
      Cuzco
      (n = 22)
      Urubamba
      (n = 22) 2010
      12.62803°−13.01153° 72.53150°−72.92381° 665−1,478 CHUN-1, CHUN-2, CHUN-3, CHUN-4, CHUN-5, CHUN-6, CHUN-7, CHUN-8, CHUN-9, CHUN-10, CHUN-11, CHUN-12, CHUN-13, CHUN-14, CHUN-15, CHUN-16, CHUN-17, CHUN-18, CHUN-19, CHUN-20, CHUN-21, CHUN-22
      Madre de Dios
      (n = 55)
      Madre de Dios
      (n = 55) 2010
      11.31142°−12.60000° 68.65986°−69.41619° 171−316 PPA-346, PPA-347, PPA-348, PPA-349, PPA-350, PPA-351, PPA-352, PPA-353, PPA-354, PPA-355, PPA-356, PPA-357, HIT-358, HIT-359, HIT-360, HIT-361, HIT-362, HIT-363, HIT-364, PPR-365, PPR-366, PPR-367, PPR-368, PPR-369, SFR-370, SFR-371, SFR-372, SFR-373, SFR-374, SFR-375, SFR-376, SFR-377, SFR-378, SND-379, SND-380, SND-381, SND-382, SND-383, SND-384, SND-385, ROL-385, ROL-386, ROL-386, ROL-387, ROL-388, ROL-389, ARR-390, ARR-391, ARR-392, ARR-393, ARR-394, ARR-395, ARR-396, ARR-397, ARR-398, ARR-399, ARR-400
      Ucayali
      (n = 34)
      Ucayali (n = 34)
      2010
      8.20719°−8.69872° 74.16783°−74.40353° 153−164 TAM-401, TAM-402, TAM-403, TAM-404, TAM-405, TAM-406, TAM-407, TAM-408, TAM-409, TAM-410, TAM-411, TAM-412, TAM-413, TAM-414, TAM-415, TAM-416, TAM-417, ABJ-418, ABJ-419, ABJ-420, ABJ-421, ABJ-422, UTQ-423, UTQ-424, UTQ-425, UTQ-426, UTQ-427, UTQ-428, UTQ-429, UTQ-430, UTQ-431, UTQ-432, UTQ-433, UTQ-434

      Table 1. 

      Accessions and location of collection of 511 wild cacao, planted and preserved at the germplasm bank of ICT in Tarapoto, Peru.

    • Descriptor
      Leaf 1 Length (LL) and width (WL) of the leaf in centimeters
      2 Length from base to the most comprehensive point of a leaf (LBL) in centimeters
      3 Petiole length (PL) in centimeters
      Fruit 4 Fruit weight (FW) in grams
      5 Seed fresh weight (SFW) and Seed dry weight (SDW)
      6 Length (FL) and diameter (FD) of fruits in centimeters
      7 Length (FL)/diameter (FD) ratio (FL/FD)
      8 Husk thickness (HusT) in centimeters
      9 Furrow depth (FwD) in centimeters
      10 Pod index (PI) [1,000/SDW]
      Seed 11 Seed number per fruit (SeN)
      12 Seed index (SI) [SDW/SeN]
      13 Length (SeL) and width (SeW) of the seeds in millimeters
      14 Seed thickness (SeT) in millimeters
      Flower 15 Length (SepL) and width (SepW) of the sepal in millimeters
      16 Length (PeL) and width (PeW) of the petal in millimeters
      17 Staminodium length (StL) in millimeters
      18 Length (OvL) and width (OvW) of the ovary in millimeters
      19 Style Length (StyL) in millimeters
      20 Number of ovules per ovary (Ov/o)

      Table 2. 

      Agro-morphological descriptors considered in this study.

    • Variables* Min Max Mean ± SD CV % F p-value
      LL (cm) 23.54 34.4 28.78 ± 3.92 13.63 1.13 0.03871
      LW (cm) 6.88 13.36 9.89 ± 2.10 21.18 1.19 0.00507
      LBL (cm) 11.52 20.52 14.65 ± 2.57 17.53 1.20 0.00329
      PL (cm) 1.26 4.94 1.81 ± 0.53 29.39 1.46 < 0.0001
      FW (g) 348.34 763.27 568.25 ± 179.64 31.61 0.95 0.7533
      SFW (g) 62.54 166.13 100.7 ± 30.55 30.34 1.42 < 0.0001
      SDW (g) 24.37 63.2 36.75 ± 10.65 28.99 1.50 < 0.0001
      FL (cm) 15.04 22.36 18.78 ± 2.05 10.94 1.68 < 0.0001
      FD (cm) 6.72 14.25 11.36 ± 2.06 18.17 3.86 < 0.0001
      FL/FD 1.24 2.9 1.71 ± 0.39 22.70 2.99 < 0.0001
      FwD (mm) 5.5 16.93 8.3 ± 2.74 33.02 2.88 < 0.0001
      HusT (mm) 10.13 21.3 15.49 ± 3.70 23.86 3.20 < 0.0001
      SeN (n) 21.2 46.8 33.09 ± 7.59 22.93 1.15 0.02259
      SI (g) 0.8 1.64 1.13 ± 0.26 23.34 1.80 < 0.0001
      PI (n) 16.4 43.6 29.67 ± 8.32 28.03 1.41 < 0.0001
      SeL (mm) 16.99 26.54 21.57 ± 2.78 12.89 3.45 < 0.0001
      SeW (mm) 10.2 15.45 12.53 ± 1.32 10.56 3.74 < 0.0001
      SeT (mm) 5.99 12.02 9.3 ± 1.58 16.95 2.25 < 0.0001
      SepL (mm) 4.88 9.5 7.21 ± 1.19 16.45 1.74 < 0.0001
      SepW (mm) 1.44 2.97 2.14 ± 0.45 21.02 1.74 < 0.0001
      PeL (mm) 3 6.44 4.52 ± 0.80 17.78 6.48 < 0.0001
      PeW (mm) 1.24 2.8 1.93 ± 0.41 21.39 4.52 < 0.0001
      StL (mm) 3.76 7.04 5.19 ± 0.68 13.16 2.86 < 0.0001
      OvL (mm) 1.16 2.28 1.68 ± 0.32 18.92 2.33 < 0.0001
      OvW (mm) 0.72 1.6 1.16 ± 0.28 23.71 2.88 < 0.0001
      StyL (mm) 1.3 2.98 2.31 ± 0.36 15.76 2.97 < 0.0001
      Ov/o (n) 22.4 47.4 35.59 ± 7.35 20.65 4.89 < 0.0001
      * Leaf traits: Leaf length (LL), Leaf width (LW), Length from base to widest point of a leaf (LBL), Petiole length (PL). Fruit traits: Fruit weight (FW), Fruit length (FL), Fruit diameter (FD), Fruit length-to-diameter ratio (FL/FD), Furrow depth (FwD), Husk thickness (HusT), Pod index (PI). Seed traits: Seed fresh weight (SFW), Seed dry weight (SDW), Seed number per fruit (SeN), Seed index (SI), Seed length (SeL), Seed width (SeW), Seed thickness (SeT). Floral traits: Sepal length (SepL), Sepal width (SepW), Petal length (PeL), Petal width (PeW), Staminate length (StL), Ovary length (OvL), Ovary width (OvW), Style length (StyL), Ovules per ovary (Ov/o). CV = coefficient of variability; F and p-value determined from analysis of variance.

      Table 3. 

      Minimum (min), maximum (max), mean values ± standard deviation (SD), and statistical significance (p ≤ 0.05) for 27 leaf, fruit, seed, and flower descriptors of 511 accessions of wild cacao from the Peruvian Amazonia.

    • Traits** River basin*
      AMZ AYP CHA MDD MAR MOR NAP NUC PAS PUT SAN TIG UCA UGU UNG URI URU YAV CV % F p-value
      LL (cm) 28.68 28.36 28.66 29.14 28.34 28.82 29.15 27.85 28.77 28.12 28.85 28.97 29.25 29.59 28.47 29.01 28.77 28.34 6.34 1.47 0.0986
      LW (cm) 9.86 b 10.38 a 10.10 a 9.62 b 9.61 b 9.71 b 10.05 a 10.10 a 10.46 a 9.94 a 9.86 b 9.73 b 9.97 a 10.19 a 9.76 b 9.64 b 9.49 b 10.33 a 9.95 2.22 0.0034
      LBL (cm) 14.36 b 15.01 a 14.71 b 14.28 b 14.34 b 14.45 b 14.66 b 15.06 a 15.31 a 14.54 b 14.47 b 14.29 b 14.58 b 15.52 a 14.73 b 14.94 a 14.18 b 14.66 b 8.22 2.53 0.0007
      PL (cm) 1.66 1.93 1.85 1.78 1.84 1.79 1.74 1.82 1.86 1.81 1.80 1.79 1.81 1.82 1.92 1.80 1.75 1.81 15.15 1.18 0.2788
      FW (g) 559.39 549.79 587.93 557.59 583.63 576.50 573.22 587.79 561.59 567.38 565.02 565.05 574.11 568.96 564.30 561.19 558.57 574.62 13.99 0.46 0.9684
      SFW (g) 110.12 a 94.07 c 105.33 b 117.48 a 99.64 b 89.6 d 100.55 b 83.52 d 90.63 d 103.26 b 103.49 b 102.56 b 105.15 b 101.53 b 101.89 b 95.92 c 95.51 c 104.47 b 13.21 12.40 <0.0001
      SDW (g) 39.88 b 34.12 c 38.26 b 42.98 a 36.20 b 32.85 d 36.62 b 30.60 d 33.17 d 37.72 b 37.60 b 37.55 b 38.27 b 36.96 b 37.33 b 35.16 c 34.80 c 38.1 b 12.81 12.95 <0.0001
      FL (cm) 19.07 a 18.93 a 17.27 b 19.11 a 18.84 a 18.65 a 19.19 a 18.85 a 19.16 a 19.08 a 16.85 b 19.01 a 19.22 a 18.65 a 19.11 a 18.96 a 19.18 a 19.30 a 4.84 16.28 <0.0001
      FD (cm) 12.15 a 12.33 a 11.78 a 11.91 a 12.06 a 8.94 b 12.03 a 11.93 a 12.04 a 12.04 a 9.21 b 12.15 a 12.01 a 12.08 a 12.01 a 12.08 a 11.66 a 8.93 b 6.74 78.11 <0.0001
      FL/FD 1.60 c 1.56 c 1.50 c 1.64 c 1.60 c 2.17 a 1.63 c 1.61 c 1.62 c 1.62 c 1.87 b 1.59 c 1.63 c 1.58 c 1.63 c 1.60 c 1.68 c 2.17 a 9.04 51.99 <0.0001
      FwD (mm) 7.31 d 8.26 c 7.46 d 7.58 d 7.98 c 11.88 a 7.88 c 8.43 c 7.93 c 7.32 d 10.30 b 7.66 d 7.23 d 8.20 c 7.83 c 7.90 c 6.62 e 7.50 d 13.16 50.06 <0.0001
      HusT (mm) 13.68 d 17.36 b 13.5 d 13.8 d 16.85 b 18.09 a 13.44 d 17.24 b 17.5 b 13.08 d 15.44 c 13.63 d 13.83 d 17.34 b 17.59 b 17.51 b 11.38 e 13.45 d 8.96 65.67 <0.0001
      SeN (n) 31.98 c 32.69 c 31.69 c 34.69 b 33.09 c 32.70 c 31.8 c 33.34 c 33.00 c 33.51 c 32.72 c 31.38 c 34.32 b 32.58 c 33.33 c 31.33 c 36.67 a 33.02 c 10.34 3.86 <0.0001
      SI (g) 1.25 a 1.05 c 1.22 a 1.25 a 1.10 b 1.01 c 1.16 b 0.92 d 1.01 c 1.15 b 1.17 b 1.22 a 1.14 b 1.15 b 1.14 b 1.15 b 0.95 d 1.19 b 10.10 20.76 <0.0001
      PI (n) 27.84 c 31.85 b 28.77 c 26.20 d 29.93 c 31.98 b 29.65 c 34.18 a 32.13 b 28.47 c 28.58 c 28.34 c 28.16 c 29.00 c 28.64 c 30.21 c 34.61 a 28.51 c 12.36 11.28 <0.0001
      SeL (mm) 20.57 c 23.94 a 20.34 c 20.41 c 23.17 a 20.09 c 20.70 c 23.69 a 23.56 a 20.53 c 21.41 b 20.64 c 20.62 c 23.54 a 23.32 a 23.63 a 18.35 d 20.41 c 4.70 75.80 <0.0001
      SeW (mm) 12.32 b 13.30 a 12.46 b 12.35 b 13.33 a 11.05 d 12.6 b 13.30 a 13.30 a 12.53 b 11.86 c 12.43 b 12.43 b 13.35 a 13.29 a 13.40 a 10.99 d 12.45 b 4.07 68.65 <0.0001
      SeT (mm) 8.98 c 9.97 a 8.74 c 9.12 c 9.67 b 9.04 c 8.79 c 10.13 a 10.05 a 9.15 c 9.08 c 8.97 c 8.93 c 10.08 a 10.09 a 10.29 a 6.74 d 8.96 c 6.23 50.91 <0.0001
      SepL (mm) 7.23 c 8.00 a 7.95 a 7.66 b 6.32 e 7.19 c 7.20 c 7.25 c 7.10 c 7.19 c 7.04 c 7.25 c 6.88 d 7.02 c 7.03 c 6.83 d 7.84 a 6.85 d 7.40 16.06 <0.0001
      SepW (mm) 2.17 b 2.73 a 2.23 b 2.21 b 2.03 c 2.27 b 2.07 c 2.29 b 2.09 c 2.07 c 1.94 d 2.02 c 2.06 c 1.98 d 2.04 c 2.06 c 2.25 b 1.98 d 8.69 24.26 <0.0001
      PeL (mm) 4.24 e 6.01 a 4.15 f 4.11 f 5.05 c 4.69 d 4.20 e 5.37 b 5.34 b 4.20 e 4.33 e 4.12 f 4.10 f 5.09 c 5.04 c 4.08 f 3.38 g 4.16 f 5.33 177.84 <0.0001
      PeW (mm) 1.79 f 2.69 a 1.70 g 1.75 f 2.00 e 2.07 d 1.79 f 2.26 c 2.06 d 1.67 g 1.97 e 1.70 g 1.72 g 2.45 b 1.93 e 1.78 f 1.66 g 1.72 g 7.95 86.33 <0.0001
      StL (mm) 5.18 d 5.78 b 5.07 e 5.29 d 5.20 d 5.01 e 5.16 d 5.49 c 5.54 c 5.22 d 4.39 f 5.18 d 5.25 d 5.00 e 5.00 e 5.11 e 5.99 a 5.23 d 5.69 37.92 <0.0001
      OvL (mm) 1.72 c 1.91 a 1.68 c 1.71 c 1.78 b 1.82 b 1.76 c 1.80 b 1.75 c 1.69 c 1.41 e 1.73 c 1.75 c 1.29 f 1.54 d 1.72 c 1.57 d 1.70 c 7.57 40.05 <0.0001
      OvW (mm) 1.29 a 1.03 d 1.27 b 1.25 b 0.89 f 1.30 a 1.33 a 0.97 e 1.03 d 1.30 a 0.98 e 1.27 b 1.29 a 1.02 d 1.00 d 1.24 b 1.11 c 1.33 a 9.15 55.53 <0.0001
      StyL (mm) 2.23 c 2.73 a 2.25 c 2.22 c 2.20 d 2.26 c 2.24 c 2.67 a 2.52 b 2.29 c 2.15 d 2.23 c 2.27 c 2.31 c 2.65 a 2.24 c 1.90 e 2.26 c 6.17 54.36 <0.0001
      Ov/o (n) 39.19 c 32.08 e 38.13 c 38.31 c 29.78 f 42.68 a 37.70 c 30.02 f 33.22 e 39.40 c 41.01 b 37.7 c 37.96 c 27.06 h 28.54 g 29.44 f 35.32 d 37.05 c 7.06 112.25 <0.0001
      * AMZ = Amazonas, AYP = Aypena, CHA = Chambira, MDD = Madre de Dios, MAR = Marañón, MOR = Morona, NAP = Napo, NUC = Nucuray, PAS = Pastaza, PUT = Putumayo, SAN = Santiago, TIG = Tigre, UCA = Ucayali, UGU = Ungumayo, UNG = Ungurahui, URI = Urituyacu, YAV = Yavarí. Means with identical letters in rows are not significantly different. ** Leaf traits: Leaf length (LL), Leaf width (LW), Length from base to the widest point of a leaf (LBL), Petiole length (PL). Fruit traits: Fruit weight (FW), Fruit length (FL), Fruit diameter (FD), Fruit length-to-diameter ratio (FL/FD), Furrow depth (FwD), Husk thickness (HusT), Pod index (PI). Seed traits: Seed fresh weight (SFW), Seed dry weight (SDW), Seed number per fruit (SeN), Seed index (SI), Seed length (SeL), Seed width (SeW), Seed thickness (SeT). Floral traits: Sepal length (SepL), Sepal width (SepW), Petal length (PeL), Petal width (PeW), Staminate length (StL), Ovary length (OvL), Ovary width (OvW), Style length (StyL), Ovules per ovary (Ov/o).

      Table 4. 

      Means for 27 leaf, fruit, seed, and flower descriptors of 511 accessions of wild cacao by river basin in peruvian amazonia. CV = coefficient of variability; F and p-value determined from analysis of variance.

    • Traits* Minimum value Maximum value
      LL (cm) AYP 10 = 23.54 UGU 114 = 34.4
      LW (cm) UNG 64 = 6.88 AYP 22 = 13.36
      LBL (cm) SAN 199 = 11.52 AYP 2 = 20.52
      PL (cm) AMZ 452 = 1.26 AYP 7 = 4.94
      FW (g) URI 165 = 348.34 TIG 307 = 763.27
      SFW (g) NAP 328 = 62.54 ARR 400 = 166.13
      SDW (g) NAP 328 = 24.37 ARR 400 = 63.2
      FL (cm) SAN 235 = 15.04 MOR 212 = 22.36
      FD (cm) MOR 220 = 6.72 PUT 476 = 14.25
      FL/FD CHA 295 = 1.24 MOR 212 = 2.9
      FwD (mm) ABI 418 = 5.5 MOR 216 = 16.93
      HusT (mm) CHUN 5 = 10.13 UNG 50 = 21.3
      SeN (n) CHA 291 = 21.2 ARR 400 = 46.8
      SI (g) CHUN 12 = 0.8 AMZ 446 = 1.64
      PI (n) ARR 400 = 16.4 YAV 459 = 43.6
      SeL (mm) CHUN 15 = 16.99 UNG 55 = 26.54
      SeW (mm) MOR 204 = 10.2 PAS 96 = 15.45
      SeT (mm) CHUN 6 = 5.99 URI 178 = 12.02
      SepL (mm) YAV 437 = 4.88 CHUN 9 = 9.5
      SepW (mm) PUT 460A, YAV 437 = 1.44 AYP 8 = 2.97
      PeL (mm) CHUN 16 = 3 AYP 1 = 6.44
      PeW (mm) TAM 415 = 1.24 AYP 9, AYP 19 = 2.8
      StL (mm) SAN 233 = 3.76 CHUN 4 = 7.04
      OvL (mm) SAN 259 = 1.16 MOR 226 = 2.28
      OvW (mm) CHUN 11 = 0.72 UTQ 423, YAV 475 = 1.6
      StyL (mm) CHUN 8 = 1.3 UNG 63 = 2.98
      Ov/o (n) UGU 129 = 22.4 MOR 248 = 47.4
      * Leaf traits: Leaf length (LL), Leaf width (LW), Length from base to the widest point of a leaf (LBL), Petiole length (PL). Fruit traits: Fruit weight (FW), Fruit length (FL), Fruit diameter (FD), Fruit length-to-diameter ratio (FL/FD), Furrow depth (FwD), Husk thickness (HusT), Pod index (PI). Seed traits: Seed fresh weight (SFW), Seed dry weight (SDW), Seed number per fruit (SeN), Seed index (SI), Seed length (SeL), Seed width (SeW), Seed thickness (SeT). Floral traits: Sepal length (SepL), Sepal width (SepW), Petal length (PeL), Petal width (PeW), Staminate length (StL), Ovary length (OvL), Ovary width (OvW), Style length (StyL), Ovules per ovary (Ov/o).

      Table 5. 

      Cacao accessions with minimum and maximum values for 27 leaf, fruit, seed, and flower descriptors.

    • River basin Phenotypic groups Total accessions by
      river basin
      Group 1 Group 2 Group 3 Group 4 Group 5
      Amazonas 17 (9.1%) 17
      Aypena 22 (11.6%) 22
      Chambira 20 (10.7%) 20
      Madre de Dios 53 (28.3%) 1 (3.8%) 1 (1.2%) 55
      Marañón 22 (11.6%) 22
      Morona 50 (58.1%) 50
      Napo 22 (11.8%) 22
      Nucuray 25 (13.2%) 25
      Pastaza 24 (12.6%) 24
      Putumayo 18 (9.6%) 18
      Santiago 1 (0.5%) 3 (11.5%) 34 (39.5%) 38
      Tigre 22 (11.8%) 22
      Ucayali 34 (18.2%) 34
      Ungumayo 26 (13.7%) 26
      Ungurahui 38 (20.0%) 38
      Urituyacu 33 (17.4%) 33
      Urubamba 22 (100%) 22
      Yavari 22 (84.6%) 1 (1.2%) 23
      Total group 187 26 190 22 86 511

      Table 6. 

      Conformation of five phenotypic groups for 511 wild cacao accessions based on river basin of origin.

    • Traits* Group 1 Group 2 Group 3 Group 4 Group 5 CV % F p-value
      LL (cm) 28.94 ± 0.13 28.39 ± 0.36 28.65 ± 0.13 28.77 ± 0.39 28.85 ± 0.2 6.39 0.95 0.4329
      LW (cm) 9.86 ± 0.07 a 10.27 ± 0.2 a 9.99 ± 0.07 a 9.49 ± 0.21 b 9.76 ± 0.11 a 10.09 2.66 0.0318
      LBL (cm) 14.46 ± 0.09 b 14.58 ± 0.24 b 14.98 ± 0.09 a 14.18 ± 0.26 b 14.47 ± 0.13 b 8.27 5.91 0.0001
      PL (cm) 1.78 ± 0.02 1.80 ± 0.05 1.86 ± 0.02 1.75 ± 0.06 1.79 ± 0.03 15.13 2.07 0.0830
      FW (g) 567.72 ± 5.78 575.01 ± 15.5 567.70 ± 5.73 558.57 ± 16.85 571.03 ± 8.52 13.91 0.16 0.9579
      SFW (g) 108.10 ± 1.06 a 104 ± 2.85 a 95.80 ± 1.05 b 95.51 ± 3.10 b 95.80 ± 1.57 b 14.42 21.03 < 0.0001
      SDW (g) 39.43 ± 0.38 a 37.79 ± 1.01 a 34.99 ± 0.37 b 34.80 ± 1.10 b 35.01 ± 0.56 b 14.06 21.55 < 0.0001
      FL (cm) 18.90 ± 0.08 a 19.06 ± 0.21 a 18.94 ± 0.08 a 19.18 ± 0.22 a 17.95 ± 0.11 b 5.61 16.43 < 0.0001
      FD (cm) 12.01 ± 0.05 a 8.92 ± 0.15 b 12.07 ± 0.05 a 11.66 ± 0.16 a 9.06 ± 0.08 b 6.60 348.70 < 0.0001
      FL/FD 1.61 ± 0.01 c 2.14 ± 0.03 a 1.60 ± 0.01 c 1.68 ± 0.04 c 2.05 ± 0.02 b 9.63 176.65 < 0.0001
      FwD (mm) 7.50 ± 0.08 c 7.45 ± 0.22 c 8.05 ± 0.08 b 6.62 ± 0.24 d 11.28 ± 0.12 a 13.43 196.41 < 0.0001
      HusT (mm) 13.63 ± 0.11 b 13.50 ± 0.29 b 17.37 ± 0.11 a 11.38 ± 0.31 c 17.03 ± 0.16 a 9.50 230.96 < 0.0001
      SeN (n) 33.17 ± 0.26 b 32.50 ± 0.69 b 32.74 ± 0.25 b 36.67 ± 0.75 a 32.96 ± 0.38 b 10.60 6.45 < 0.0001
      SI (g) 1.20 ± 0.01 a 1.20 ± 0.03 a 1.08 ± 0.01 b 0.95 ± 0.03 c 1.07 ± 0.01 b 11.48 37.4 < 0.0001
      PI (n) 27.86 ± 0.29 c 28.74 ± 0.77 c 30.65 ± 0.28 b 34.61 ± 0.84 a 30.45 ± 0.42 b 13.23 22.78 < 0.0001
      SeL (mm) 20.53 ± 0.08 b 20.36 ± 0.21 b 23.54 ± 0.08 a 18.35 ± 0.22 c 20.67 ± 0.11 b 4.85 290.59 < 0.0001
      SeW (mm) 12.43 ± 0.04 b 12.39 ± 0.1 b 13.33 ± 0.04 a 10.99 ± 0.11 d 11.39 ± 0.06 c 4.26 251.82 < 0.0001
      SeT (mm) 8.97 ± 0.04 b 8.95 ± 0.12 b 10.06 ± 0.04 a 6.74 ± 0.13 c 9.08 ± 0.06 b 6.31 204.54 < 0.0001
      SepL (mm) 7.36 ± 0.05 b 6.85 ± 0.12 c 7.06 ± 0.05 c 7.84 ± 0.13 a 7.14 ± 0.07 c 8.65 13.60 < 0.0001
      SepW (mm) 2.13 ± 0.02 a 2.00 ± 0.05 b 2.15 ± 0.02 a 2.25 ± 0.05 a 2.13 ± 0.03 a 11.48 3.41 0.0092
      PeL (mm) 4.14 ± 0.03 c 4.20 ± 0.08 c 5.07 ± 0.03 a 3.38 ± 0.09 d 4.53 ± 0.05 b 9.24 165.64 < 0.0001
      PeW (mm) 1.74 ± 0.02 c 1.74 ± 0.05 c 2.13 ± 0.02 a 1.66 ± 0.05 c 2.02 ± 0.03 b 12.29 78.65 < 0.0001
      StL (mm) 5.21 ± 0.03 b 5.08 ± 0.07 b 5.26 ± 0.03 b 5.99 ± 0.08 a 4.79 ± 0.04 c 7.15 53.71 < 0.0001
      OvL (mm) 1.72 ± 0.01 a 1.66 ± 0.04 b 1.67 ± 0.01 b 1.57 ± 0.04 b 1.66 ± 0.02 b 11.33 4.55 0.0013
      OvW (mm) 1.28 ± 0.01 a 1.27 ± 0.03 a 1.04 ± 0.01 d 1.11 ± 0.03 c 1.17 ± 0.02 b 12.26 73.40 < 0.0001
      StyL (mm) 2.25 ± 0.01 b 2.26 ± 0.04 b 2.48 ± 0.01 a 1.90 ± 0.04 c 2.21 ± 0.02 b 8.23 72.96 < 0.0001
      Ov/o (n) 38.23 ± 0.20 b 37.61 ± 0.54 b 29.83 ± 0.20 d 35.32 ± 0.59 c 42.01 ± 0.30 a 7.80 364.99 < 0.0001
      * Leaf traits: Leaf length (LL), Leaf width (LW), Length from base to the widest point of a leaf (LBL), Petiole length (PL). Fruit traits: Fruit weight (FW), Fruit length (FL), Fruit diameter (FD), Fruit length-to-diameter ratio (FL/FD), Furrow depth (FwD), Husk thickness (HusT), Pod index (PI). Seed traits: Seed fresh weight (SFW), Seed dry weight (SDW), Seed number per fruit (SeN), Seed index (SI), Seed length (SeL), Seed width (SeW), Seed thickness (SeT). Floral traits: Sepal length (SepL), Sepal width (SepW), Petal length (PeL), Petal width (PeW), Staminate length (StL), Ovary length (OvL), Ovary width (OvW), Style length (StyL), Ovules per ovary (Ov/o). Means with the same letter in a row are not statistically significant (p > 0.05).

      Table 7. 

      Mean values ± standard error and statistical significance for 27 leaf, fruit, seed, and flower descriptors for five cacao phenotypic groups from Peruvian Amazonia.